Thesis (PhD)--Stellenbosch University, 2014. / ENGLISH ABSTRACT: Soft, serene insect songs add an intrinsic aesthetic value to the landscape. Yet these songs also have an important biological relevance. Acoustic signals across the landscape carry a multitude of localized information allowing organisms to communicate invisibly within their environment. Ensifera are cryptic participants of nocturnal soundscapes, contributing to ambient acoustics through their diverse range of proclamation songs. Although not without inherent risks and constraints, the single most important function of signalling is sexual advertising and pair formation. In order for acoustic communication to be effective, signals must maintain their encoded information so as to lead to positive phonotaxis in the receiver towards the emitter. In any given environment, communication is constrained by various local abiotic and biotic factors, resulting in Ensifera utilizing acoustic niches, shifting species songs spectrally, spatially and temporally for their optimal propagation in the environment. Besides the importance of Ensifera songs from an ethological point of view, the multitude of species-specific signals provide an acoustic tapestry representing species diversity across ecological gradients and over time. Acoustic inventorying and monitoring of the landscape can reflect the environmental status of ecological systems, from natural to disturbed by human influence. In contrast to traditional survey techniques, sound recording and interpretation is a non-invasive method that allows for the detection and classification of highly cryptic, yet insightful indicators of ecosystem change. Here, acoustic monitoring was used across diverse ecological gradients to improve understanding of species diversity patterns, and how they change in response to both natural gradients and in response to the human dominated landscape mosaic. This study was undertaken in three geographic locations from tropical rain forest of Brunei, Borneo, to the landscapemosaic of plantation forestry in KwaZulu-Natal, South Africa, and to the botanically rich, mountain fynbos region of the Cape Floristic Region, also in South Africa. Each region provided a diverse and particular landscape to test the value of acoustic surveys for determining local diversity patterns across natural gradients and to assess the value of the technique for assessing the impact human influence across landscapes. In tropical rainforests, an entire acoustic guild was investigated to determine how acoustic species partition their acoustic communication channels spectrally, temporally and spatially, to avoid acoustic interference. The overall assemblage showed considerable spectral partitioning. Diurnally active species showed low temporal niche overlap, whereas nocturnal species did not utilize temporal partitioning. Lack of nocturnal temporal partitioning suggests other mechanisms of acoustic avoidance are sufficient to avoid acoustic overlap, or that there are insufficient cues to partition nocturnal acoustic environments. Acoustic species also utilized spatial partitioning, with distinct acoustic assemblages at vertical heights and with elevation. Utilization of a range of different strategies allow many species to communicate with conspecifics with little or no interference from other species in a signal rich environment. Acoustic profiling was also undertaken in KwaZulu-Natal, South Africa, across a plantation forestry landscape mosaic with diverse ecological gradients containing both alien and indigenous vegetation, as well as boarding large natural protected areas. Areas covered in alien timber or non-endemic grass were devoid of acoustic signals. Managed areas that were mowed and heavily grazed were not effective in maintaining the natural complement of nocturnal acoustic species. Within natural vegetation patches inside plantations, acoustic species richness increased with plant heterogeneity and patch size. Patches of indigenous vegetation within the plantation matrix effectively reduced the contrast of transformed landscapes with surrounding natural areas, with indigenous forest patches containing a highly characteristic acoustic species assemblage. Within the botanically rich, mountain fynbos region of the Cape Floristic Region, acoustic profiling was conducted across gradients of elevation, season and vegetation. Across these gradients, katydid acoustic signals were identified and characterized for the first time. This resulted in the discovery of two new katydid species and a novel sound producing structure in a carabid beetle, a species previously unknown to produce sound. Acoustic diversity across seasonal and elevational gradients increased with increasing temperatures. Climatic variability along the elevational gradient produced variation in seasonal phenology. Katydids also utilized high frequency acoustic signals, which is probably an adaptation to overcome background noise from wind, so prevalent in this area. Furthermore, despite producing conspicuous signals for mate attraction and pair formation, katydids were found not to be part of bat-eared fox diet, an insectivorous, nocturnal predator that uses its characteristic large ears to detect sounds made by invertebrate prey. This study shows the value of using acoustic emissions from katydids to identify acoustic diversity patterns across ecological gradients and in response to human impacts on the landscape. / AFRIKAANSE OPSOMMING: Sagte, rustige insekliedjies voeg ‘n intrinsieke estetiese waarde aan die landskap. Tog is hierdie liedjies ook van belangrike biologiese waarde. Akoestiese seine oor die landskap dra ‘n magdom plaaslike inligting wat organismes in staat stel om onsigbaar te kommunikeer binne hul omgewing. Langhoringsprinkane is kriptiese deelnemers van die nagtelike klankomgewing en dra by tot die omringende akoestiek deur hul verskeidenheid van proklamasieliedjies. Alhoewel dit nie sonder inherente risiko’s en beperkings is nie, is die belangrikste funksie van seine seksuele advertering en paarvorming. Vir akoestiese kommunikasie om effektief te wees, moet seine hul geënkodeerde inligting handhaaf, sodat dit sal lei tot positiewe fonotaksis in die ontvanger teenoor die emittor. In enige gegewe omgewing, word kommunikasie beperk deur verskeie plaaslike abiotiese en biotiese faktore. Dit lei tot die gebruik van akoestiese nisse deur langhoringsprinkane, wat hulle liedjies spektraal, ruimtelik en temporeel aanpas vir optimale verspreiding in die omgewing. Benewens die belang van die langhoringsprinkaan liedjies uit ʼn etologiese oogpunt, bied die menigte spesie-spesifieke seine ʼn akoestiese tapisserie wat spesiesdiversiteit verteenwoordig oor ekologiese gradiënte en oor tyd. Akoestiese opname en monitering van die landskap kan die omgewingstoestand van ekologiese stelsels weerspieël, van natuurlike tot menslik versteurde stelsels. In teenstelling met tradisionele opnametegnieke, is klankopname en interpretasie ʼn nie-indringende metode wat dit moontlik maak om hoogs kriptiese, nog insiggewende indikators van ekosisteemverandering op te spoor en te klassifiseer. In hierdie studie is akoestiese monitering gebruik oor diverse ekologiese gradiënte om ons begrip te verbeter van spesies diversiteitspatrone, en hoe dit verander in reaksie op beide natuurlike gradiënte en in reaksie op die menslik gedomineerde landskapmosaïek. Hierdie studie is onderneem in drie geografiese liggings: tropiese reënwoud in Brunei, Borneo, die landskapmosaïek van plantasiebosbou in KwaZulu-Natal, Suid-Afrika, en die plantryke, bergfynbos-streek van die Kaap Floristiese Streek, ook in Suid-Afrika. Elke streek het 'n diverse en besondere landskap verskaf om die waarde van akoestiese opnames te toets vir die bepaling van plaaslike diversiteitspatrone in natuurlike gradiënte, asook om die waarde van die tegniek te bepaal vir die beoordeling van die impak van menslike invloed oor landskappe. In tropiese reënwoude, is 'n hele akoestiese gilde ondersoek om te bepaal hoe akoestiese spesies hul akoestiese kommunikasiekanale spektraal, ruimtelik en temporeel verdeel om akoestiese inmenging te vermy. Die algehele groep het aansienlike spektrale verdeling getoon. Dagaktiewe spesies het lae temporele nisoorvleueling getoon, terwyl nagtelike spesie nie temporele verdeling benut het nie. Gebrek aan nagtelike temporele verdeling dui daarop dat ander meganismes van akoestiese vermyding voldoende is om akoestiese oorvleueling te vermy, of dat daar onvoldoende seine is om nagtelike akoestiese omgewings te verdeel. Akoestiese spesies het ook ruimtelike verdeling benut, met verskillende akoestiese spesiesversamelings op vertikale hoogtes en met hoogte bo seevlak. Die gebruik van 'n verskeidenheid van strategieë maak dit moontlik vir spesies om te kommunikeer met min of geen inmenging van ander spesies in 'n seinryke omgewing. Akoestiese profielsamestelling is ook onderneem in KwaZulu-Natal, Suid-Afrika, oor 'n plantasiebosbou landskapmosaïek met diverse ekologiese gradiënte wat beide uitheemse en inheemse plantegroei, sowel as groot, natuurlike, beskermde gebiede ingesluit het. Gebiede wat bestaan het uit uitheemse timmerhoutbome of nie-endemiese gras, was heeltemal sonder akoestiese seine. Bestuursgebiede wat gesny en swaar bewei was, het nie doeltreffend die natuurlike komplement van nagtelike akoestiese spesies gehandhaaf nie. In natuurlike plantegroei fragmente binne plantasies, het akoestiese spesiesrykheid toegeneem met plantverskeidenheid en fragmentgrootte. Fragmente van inheemse plantegroei binne die plantasiematriks het effektief die kontras van getransformeerde landskappe met omliggende natuurlike gebiede verminder, en inheemse woudefragmente het hoogs kenmerkende akoestiese spesiesversamelings gehad. Binne die plantryke bergfynbosstreek van die Kaap Floristiese Streek, was akoestiese profielsamestelling gedoen oor gradiënte van hoogte bo seevlak, seisoen en plantegroei. Oor hierdie gradiënte, is langhoringsprinkaan akoestiese seine geïdentifiseer en gekenmerk vir die eerste keer. Dit het gelei tot die ontdekking van twee nuwe langhoringsprinkaan spesies en 'n nuwe klankvervaardiging struktuur in 'n Carabid kewer, 'n spesie wat nie voorheen bekend was om klank te produseer nie. Akoestiese diversiteit oor gradiënte van seisoen en hoogte bo seevlak het toegeneem met toenemende temperature. Variasie in klimaatstoestande oor die gradiënt van hoogte bo seevlak het variasie in seisoenale fenologie veroorsaak. Langhoringsprinkane het ook hoë frekwensie akoestiese seine benut, wat waarskynlik 'n aanpassing is om agtergrondgeraas van die wind, wat so algemeen is in hierdie gebied, te bowe te kom. Verder, ten spyte van die vervaardiging van opvallende seine vir maataantrekking en paarvorming, het langhoringsprinkane nie deel gevorm van die bakoorjakkals se dieet nie. Hierdie is 'n insekvretende, nagtelike roofdier wat gebruik maak van sy kenmerkende groot ore om klanke op te spoor wat gemaak word deur invertebraatprooi. Hierdie studie toon die waarde van die gebruik van akoestiese seine van langhoringsprinkane om akoestiese diversiteitspatrone te identifiseer oor ekologiese gradiënte en in reaksie op menslike impakte op die landskap.

Wetlands contain both aquatic and terrestrial environments which generates high biodiversity. However, they are commonly associated with mosquitoes (Diptera: Culicidae), and mosquitoes are usually regarded as negative by humans because they can cause nuisance and transmit diseases. This thesis aimed to clarify the association between mosquitoes and wetlands and to achieve a more balanced view of biodiversity in wetlands by including mosquito diversity. Studies on adult mosquito diversity and assemblages were performed in 18 wetlands spread over Sweden. The Swedish mosquito species were organized in ten functional groups based on four life-history characteristics. This classification was used as an additional diversity measurement and as a tool for presentation of mosquito data. Mosquito diversity showed several of the well-established diversity patterns such as a latitudinal gradient, a species-area relationship and a distribution-abundance relationship. In a landscape perspective, diversity of both mosquitoes and dytiscids (Coleoptera: Dytiscidae) were positively influenced by a high proportion of permanent water and a high amount of open areas, indicating co-varying diversity patterns. Mosquito assemblages in the Nedre Dalälven region were mainly structured by the extent of flooded areas and wetland type (wet meadow, swamp and bog). In addition to the influence of the proportion of temporary wetlands at a local scale, the proportion of forest gained importance at larger spatial scales and in relation to dispersal distances of species. In southern Sweden, mosquito faunas differed between natural and constructed wetlands, partly reflecting differences in wetland size. In an experiment, different responses of two co-occurring mosquito species to rapid larval habitat desiccation indicate that weather conditions after a flood could influence mosquito assemblages. The conclusions of this thesis provide suggestions on how to construct and position wetlands for increased insect diversity, and indicate that low abundance of major nuisance species might be crucial for acceptance of wetlands near human settlements.

No description available.

L'objectif de cette thèse a été de caractériser les patrons spatiaux de diversité inter- et intraspécifique au sein des réseaux dendritiques, d'expliciter les processus évolutifs et écologiques qui les sous-tendent, et d'isoler les possibles covariations spatiales et interactions existant entre ces différentes facettes de biodiversité. Pour cela, j'ai tout d'abord développé de nouvelles méthodes statistiques permettant l'analyse, par des modèles causaux, de données sous la forme de matrices de distances, afin de pouvoir analyser plusieurs facettes de biodiversité dans un unique cadre statistique au niveau alpha et bêta. J'ai par la suite étudié de manière intégrative les patrons de diversité interspécifique et intraspécifique génétique d'une part, et intraspécifique génétique et intraspécifique phénotypique d'autre part, au sein du bassin versant Garonne- Dordogne. Enfin, j'ai utilisé un modèle de dynamique éco-évolutive afin d'étudier l'impact de la structure et des gradients environnementaux caractérisant les réseaux dendritiques sur l'adaptation locale au sein de ces réseaux. / The aim of this thesis was to characterized the spatial patterns of inter- and intraspecific diversity within riverine networks, to better understand the ecological and evolutionary processes underlying them and to explore how the different facets of biodiversity interact with one another. First, I developed novel statistical approaches allowing the application of causal modeling to data in the form of pairwise matrices, thus allowing the study within integrative frameworks of several biodiversity facets at the alpha and beta levels. I then studied integratively the patterns of interspecific and intraspecific genetic diversity and of intraspecific genetic and intraspecific phenotypic diversity within the Garonne-Dordogne river basin. Finally, I used an eco-evolutionary metapopulation dynamics model to assess the impacts of the structure and environmental gradients that characterize riverine networks on local adaptation.

Les patrons de diversité caractérisent la structure de la diversité des communautés, c'est-à-dire sa valeur, sa distribution et son changement dans l'espace et le temps. Leur étude peut amener des informations importantes sur les processus écologiques qui en sont à l'origine. Cependant de nombreuses hypothèses de travail sont faites lors de leur analyse. L'idée générale de cette thèse est qu'en remettant en cause ces hypothèses, un certain nombre de développements liés aux indices de diversité et aux modèles nuls deviennent possibles et permettent de mieux comprendre les processus écologiques à l'origine des patrons de diversité fonctionnelle ou phylogénétique. Le premier chapitre est consacré à l'étude des patrons de diversité fonctionnelle des communautés végétales alpines à de multiples échelles spatiales et organisationnelles. Le second chapitre s'intéresse aux perspectives méthodologiques amenés par les nombres de Hill. Dans le dernier chapitre, on s'intéresse aux enjeux méthodologiques d'un nouveau type de données de communautés : l'ADN environnemental. / Patterns of community diversity refers to the structure of diversity, i.e. its quantification, its distribution and its turnover in space and time. Its study is likely to shed the light on the assembly rules that determined the structure of communities. However, numerous ecological assumptions are often made when studying diversity patterns. What motivated the work was the perspective that by relaxing these assumptions, a number of developments linked to diversity indices and null models are possible and can help to understand the impact of multiple ecological processes on phylogenetic and functional diversity patterns. In a first part we studied the pattern of functional diversity of alpine plant communities as a function of spatial and organizational scales. In the second part, we studied the methodological perspectives brought by the Hill numbers. In a third part, we addressed the main methodological issues of a new type of community data: environmental DNA.

Formações rupestres representam centros de diversidade taxonômica e funcional associadas a condições ambientais extremas. São também sítios para colonização por espécies florestais, normalmente associada a mecanismos de facilitação. A distribuição das espécies, determinada por seu nicho e por mecanismos neutros, resulta na variação da composição entre comunidades (diversidade beta). Neste trabalho pretendemos avaliar se a vegetação rupestre apresenta-se, taxonômica e funcionalmente, distinta da matriz florestal regional, analisar possíveis interações positivas entre espécies associadas ao avanço da floresta sobre a vegetação rupestre e determinar como a composição de espécies arbóreas no gradiente floresta-afloramento rochoso é limitada por dinâmicas neutras, de nicho ou ambas ao longo da ontogenia. Estudamos a vegetação sobre afloramentos basálticos no Parque Estadual do Turvo, sul do Brasil. A flora das comunidades rupestres foi estudada a campo durante dois anos. As espécies foram classificadas em formas de vida com base na literatura e observação no campo. Para avaliarmos a ocorrência de facilitação e os padrões de diversidade beta arbórea amostramos comunidades de espécies arbóreas em três ecótonos de floresta/afloramento rochoso. Em cada sítio estabelecemos 60 parcelas de 1.5 x 1.5 m ao longo do gradiente. As parcelas foram descritas pela cobertura de Bromelia balansae Mez e pela composição de espécies, separadas em quatro classes de tamanho, representando as diferentes fases ontogenéticas dos indivíduos. Avaliamos diferenças taxonômicas e funcionais entre a vegetação rupestre e a flora regional através de testes de qui-quadrado e os efeitos dos diferentes fatores através de testes de aleatorização uni e multivariados. Registramos 111 espécies, das quais 43 de ocorrência restrita aos afloramentos, distribuídas em 54 famílias botânicas. O espectro de formas de vida teve grande frequência de caméfitos (29.7%), nanofanerófitos (17.1%) e geófitos (15.3%). Entre as espécies de ocorrência restrita, geófitos (32.6%), caméfitos (25.6%) e nanofanerófitos (20.9%) como foram as mais frequentes. Testes de 2 indicaram diferenças significativas nas frequências de famílias e de formas de vida entre os ambientes. Nossos resultados indicaram uma associação positiva entre a cobertura de bromélias e a riqueza e a abundância de plântulas de espécies arbóreas. Aparentemente, maiores coberturas por B. balansae fornecem proteção contra a herbivoria, indistintamente, e amenização das severas condições ambientais, favorecendo o estabelecimento de espécies pioneiras da floresta. Os testes multivariados indicaram efeito significativo do fator sítio sobre a diversidade beta, em todas as classes de tamanho, e do fator gradiente somente para duas classes de tamanho analisadas. A vegetação rupestre apresentou uma flora distinta, florística e funcionalmente, com uma grande similaridade a outras formações rupestres neotropicais, mas com expressivo número de espécies arbóreas. Plântulas destas espécies, aparentemente, beneficiam-se da presença de B. balansae, o que pode levar à redução gradual da vegetação rupestre neste mosaico floresta/afloramentos. Entender como as comunidades arbóreas têm sua distribuição determinada por limitações de dispersão e de nicho pode levar a uma melhor compreensão dos processos que geram e mantêm a diversidade. / Rocky outcrops are centers of taxonomic and functional diversity with extreme environmental conditions. They also are sites for colonization by forestry species, usually related to facilitation mechanisms. The species distribution set by niche and neutrals mechanisms results in the variation of composition among communities (beta diversity). The goals of this work are evaluate if rocky outcrops are taxonomically and functionally different from the surrounding forest, study the possible positive interactions among species related to the advance of the forest over the rupestrian vegetation and determine how the composition of arboreal species in the interface forest/rocky outcrop is limited by neutral, niche or both dynamics along the ontogeny. We studied the vegetation of basaltic rocky outcrops at the Turvo State Park, south Brazil. The flora from rupestrian communities was analyzed at field during two years. Species were classified into life forms based on literature and field observations. To evaluate the patterns of arboreal beta diversity and the occurrence of facilitation between plants we sampled arboreal species in three forest/rocky outcrop ecotones. In each rocky outcrop we established 60 plots of 1.5 x 1.5 m along the forest-rocky outcrop ecotone. We described each plot in terms of Bromelia balansae Mez cover and the composition of tree species, separated into four size classes, representing different ontogenetic stages of individuals. The taxonomic and functional differences between the rupestrian vegetation and regional flora were analyzed with the chi-square test and the effects of different factors were analyzed with randomization tests uni-and multivariate. We registered 111 species, 43 of which restricted to the rocky outcrops, distributed in 54 botanic families. The life form spectrum showed a high proportion of chamaephytes (29.7%), nanophanerophytes (17.1%) and geophytes (15.3%). The flora restricted to rocky outcrops showed more frequently geophytes (32.6%), chamaephytes (25.6%) and nanophanerophytes (20.9%). Chi-square tests indicated significant differences in frequencies among environments, for both families and life forms. Our results showed that the coverage of bromeliads is positively correlated to the richness and abundance of seedlings of arboreal species. It seems that higher cover by B. balansae gives protection against herbivory indistinctively, and also reduces the effects of severe environmental conditions, allowing the settlement of forest pioneers species. Multivariate tests showed significant effect of the site factor on the beta diversity in all classes of size and significant effect of the gradient factor only on two size classes evaluated. The rupestrian vegetation is distinct floristically and functionally, showing high floristic similarity with other neotropical rupestrian formations, but has high number of arboreal species. Seedlings of arboreal species seem to have benefit from the presence of B. balansae, which can cause a gradual reduction of the rupestrian vegetation in the forest and rocky outcrop mosaic. Understanding how the distribution of arboreal communities is determined by limitations of dispersal and niche can improve the comprehension of the dynamics that generate and maintain diversity.

Formações rupestres representam centros de diversidade taxonômica e funcional associadas a condições ambientais extremas. São também sítios para colonização por espécies florestais, normalmente associada a mecanismos de facilitação. A distribuição das espécies, determinada por seu nicho e por mecanismos neutros, resulta na variação da composição entre comunidades (diversidade beta). Neste trabalho pretendemos avaliar se a vegetação rupestre apresenta-se, taxonômica e funcionalmente, distinta da matriz florestal regional, analisar possíveis interações positivas entre espécies associadas ao avanço da floresta sobre a vegetação rupestre e determinar como a composição de espécies arbóreas no gradiente floresta-afloramento rochoso é limitada por dinâmicas neutras, de nicho ou ambas ao longo da ontogenia. Estudamos a vegetação sobre afloramentos basálticos no Parque Estadual do Turvo, sul do Brasil. A flora das comunidades rupestres foi estudada a campo durante dois anos. As espécies foram classificadas em formas de vida com base na literatura e observação no campo. Para avaliarmos a ocorrência de facilitação e os padrões de diversidade beta arbórea amostramos comunidades de espécies arbóreas em três ecótonos de floresta/afloramento rochoso. Em cada sítio estabelecemos 60 parcelas de 1.5 x 1.5 m ao longo do gradiente. As parcelas foram descritas pela cobertura de Bromelia balansae Mez e pela composição de espécies, separadas em quatro classes de tamanho, representando as diferentes fases ontogenéticas dos indivíduos. Avaliamos diferenças taxonômicas e funcionais entre a vegetação rupestre e a flora regional através de testes de qui-quadrado e os efeitos dos diferentes fatores através de testes de aleatorização uni e multivariados. Registramos 111 espécies, das quais 43 de ocorrência restrita aos afloramentos, distribuídas em 54 famílias botânicas. O espectro de formas de vida teve grande frequência de caméfitos (29.7%), nanofanerófitos (17.1%) e geófitos (15.3%). Entre as espécies de ocorrência restrita, geófitos (32.6%), caméfitos (25.6%) e nanofanerófitos (20.9%) como foram as mais frequentes. Testes de 2 indicaram diferenças significativas nas frequências de famílias e de formas de vida entre os ambientes. Nossos resultados indicaram uma associação positiva entre a cobertura de bromélias e a riqueza e a abundância de plântulas de espécies arbóreas. Aparentemente, maiores coberturas por B. balansae fornecem proteção contra a herbivoria, indistintamente, e amenização das severas condições ambientais, favorecendo o estabelecimento de espécies pioneiras da floresta. Os testes multivariados indicaram efeito significativo do fator sítio sobre a diversidade beta, em todas as classes de tamanho, e do fator gradiente somente para duas classes de tamanho analisadas. A vegetação rupestre apresentou uma flora distinta, florística e funcionalmente, com uma grande similaridade a outras formações rupestres neotropicais, mas com expressivo número de espécies arbóreas. Plântulas destas espécies, aparentemente, beneficiam-se da presença de B. balansae, o que pode levar à redução gradual da vegetação rupestre neste mosaico floresta/afloramentos. Entender como as comunidades arbóreas têm sua distribuição determinada por limitações de dispersão e de nicho pode levar a uma melhor compreensão dos processos que geram e mantêm a diversidade. / Rocky outcrops are centers of taxonomic and functional diversity with extreme environmental conditions. They also are sites for colonization by forestry species, usually related to facilitation mechanisms. The species distribution set by niche and neutrals mechanisms results in the variation of composition among communities (beta diversity). The goals of this work are evaluate if rocky outcrops are taxonomically and functionally different from the surrounding forest, study the possible positive interactions among species related to the advance of the forest over the rupestrian vegetation and determine how the composition of arboreal species in the interface forest/rocky outcrop is limited by neutral, niche or both dynamics along the ontogeny. We studied the vegetation of basaltic rocky outcrops at the Turvo State Park, south Brazil. The flora from rupestrian communities was analyzed at field during two years. Species were classified into life forms based on literature and field observations. To evaluate the patterns of arboreal beta diversity and the occurrence of facilitation between plants we sampled arboreal species in three forest/rocky outcrop ecotones. In each rocky outcrop we established 60 plots of 1.5 x 1.5 m along the forest-rocky outcrop ecotone. We described each plot in terms of Bromelia balansae Mez cover and the composition of tree species, separated into four size classes, representing different ontogenetic stages of individuals. The taxonomic and functional differences between the rupestrian vegetation and regional flora were analyzed with the chi-square test and the effects of different factors were analyzed with randomization tests uni-and multivariate. We registered 111 species, 43 of which restricted to the rocky outcrops, distributed in 54 botanic families. The life form spectrum showed a high proportion of chamaephytes (29.7%), nanophanerophytes (17.1%) and geophytes (15.3%). The flora restricted to rocky outcrops showed more frequently geophytes (32.6%), chamaephytes (25.6%) and nanophanerophytes (20.9%). Chi-square tests indicated significant differences in frequencies among environments, for both families and life forms. Our results showed that the coverage of bromeliads is positively correlated to the richness and abundance of seedlings of arboreal species. It seems that higher cover by B. balansae gives protection against herbivory indistinctively, and also reduces the effects of severe environmental conditions, allowing the settlement of forest pioneers species. Multivariate tests showed significant effect of the site factor on the beta diversity in all classes of size and significant effect of the gradient factor only on two size classes evaluated. The rupestrian vegetation is distinct floristically and functionally, showing high floristic similarity with other neotropical rupestrian formations, but has high number of arboreal species. Seedlings of arboreal species seem to have benefit from the presence of B. balansae, which can cause a gradual reduction of the rupestrian vegetation in the forest and rocky outcrop mosaic. Understanding how the distribution of arboreal communities is determined by limitations of dispersal and niche can improve the comprehension of the dynamics that generate and maintain diversity.

Biodiversity is determined by a myriad of complex processes acting at different scales. Given the current rates of biodiversity loss and change, it is of paramount importance that we improve our understanding of the underlying structure of ecological communities. In this thesis, I focused on Species Abundance Distributions (SAD), as a synthetic measure of biodiversity and community structure, and on Beta (β) diversity patterns, as a description of the spatial variation of species composition. I systematically assessed the effect of scale on both these patterns, analysing a broad range of community data, including different taxa and habitats, from the terrestrial, marine and freshwater realms. Knowledge of the scaling properties of abundance and compositional patterns must be fully integrated in biodiversity research if we are to understand biodiversity and the processes underpinning it, from local to global scales. SADs depict the relative abundance of the species present in a community. Although typically described by unimodal logseries or lognormal distributions, empirical SADs can also exhibit multiple modes. However, the existence of multiple modes in SADs has largely been overlooked, assumed to be due to sampling errors or a rare pattern. Thus, we do not know how prevalent multimodality is, nor do we have an understanding of the factors leading to this pattern. Here, I provided the first global empirical assessment of the prevalence of multimodality across a wide range of taxa, habitats and spatial extents. I employed an improved method combining two model selection tools, and (conservatively) estimated that ~15% of the communities were multimodal with strong support. Furthermore, I showed that the pattern is more common for communities at broader spatial scales and with greater taxonomic diversity (i.e. more phylogenetically diverse communities, since taxonomic diversity was measured as number of families). This suggests a link between multimodality and ecological heterogeneity, broadly defined to incorporate the spatial, environmental, taxonomic and functional variability of ecological systems. Empirical understanding of how spatial scale affects SAD shape is still lacking. Here, I established a gradient in spatial scale spanning several orders of magnitude by decomposing the total extent of several datasets into smaller subsets. I performed an exploratory analysis of how SAD shape is affected by area sampled, species richness, total abundance and taxonomic diversity. Clear shifts in SAD shape can provide information about relevant ecological and spatial mechanisms affecting community structure. There was a clear effect of area, species richness and taxonomic diversity in determining SAD shape, while total abundance did not exhibit any directional effect. The results supported the findings of the previous analysis, with a higher prevalence of multimodal SADs for larger areas and for more taxonomically diverse communities, while also suggesting that species spatial aggregation patterns can be linked to SAD shape. On the other hand, there was a systematic departure from the predictions of two important macroecological theories for SAD across scales, specifically regarding logseries distributions being selected only for smaller scales and when species richness and number of families were proportionally much smaller than the total extent. β diversity quantifies the variation in species composition between sites. Although a fundamental component of biodiversity, its spatial scaling properties are still poorly understood. Here, I tested if two conceptual types of β diversity showed systematic variation with scale, while also explicitly accounting for the two β diversity components, turnover and nestedness (species replacement vs species richness differences). I provided the first empirical analysis of β diversity scaling patterns for different taxa, revealing remarkably consistent scaling curves. Total β diversity and turnover exhibit a power law decay with log area, while nestedness is largely insensitive to scale changes. For the distance decay of similarity analysis, while area sampled affected the overall dissimilarity values, rates of similarity were consistent across large variations in sampled area. Finally, in both these analyses, turnover was the main contributor to compositional change. These results suggest that species are spatially aggregated across spatial scales (from local to regional scales), while also illustrating that substantial change in community structure might occur, despite species richness remaining relatively stable. This systematic and comprehensive analysis of SAD and community similarity patterns highlighted spatial scale, ecological heterogeneity and species spatial aggregation patterns as critical components underlying the results found. This work expanded the range of scales at which both theories deriving SAD and community similarity studies have been developed and tested (from local plots to continents). The results here showed strong departures from two important macroecological theories for SAD at different scales. In addition, the overall findings in this thesis clearly indicate that unified theories of biodiversity (or assuming a set of synthetic minimal assumptions) are unable to accommodate the variability in SADs shape across spatial scales reported here, and cannot fully reproduce community similarity patterns across scales. Incorporating more realistic assumptions, or imposing scale dependent assumptions, may prove to be a fruitful avenue for ecological research regarding the scaling properties of SAD and community similarity patterns. This will allow deriving new predictions and improving the ability of theoretical models to incorporate the variability in abundance and similarity patterns across scales.

Formações rupestres representam centros de diversidade taxonômica e funcional associadas a condições ambientais extremas. São também sítios para colonização por espécies florestais, normalmente associada a mecanismos de facilitação. A distribuição das espécies, determinada por seu nicho e por mecanismos neutros, resulta na variação da composição entre comunidades (diversidade beta). Neste trabalho pretendemos avaliar se a vegetação rupestre apresenta-se, taxonômica e funcionalmente, distinta da matriz florestal regional, analisar possíveis interações positivas entre espécies associadas ao avanço da floresta sobre a vegetação rupestre e determinar como a composição de espécies arbóreas no gradiente floresta-afloramento rochoso é limitada por dinâmicas neutras, de nicho ou ambas ao longo da ontogenia. Estudamos a vegetação sobre afloramentos basálticos no Parque Estadual do Turvo, sul do Brasil. A flora das comunidades rupestres foi estudada a campo durante dois anos. As espécies foram classificadas em formas de vida com base na literatura e observação no campo. Para avaliarmos a ocorrência de facilitação e os padrões de diversidade beta arbórea amostramos comunidades de espécies arbóreas em três ecótonos de floresta/afloramento rochoso. Em cada sítio estabelecemos 60 parcelas de 1.5 x 1.5 m ao longo do gradiente. As parcelas foram descritas pela cobertura de Bromelia balansae Mez e pela composição de espécies, separadas em quatro classes de tamanho, representando as diferentes fases ontogenéticas dos indivíduos. Avaliamos diferenças taxonômicas e funcionais entre a vegetação rupestre e a flora regional através de testes de qui-quadrado e os efeitos dos diferentes fatores através de testes de aleatorização uni e multivariados. Registramos 111 espécies, das quais 43 de ocorrência restrita aos afloramentos, distribuídas em 54 famílias botânicas. O espectro de formas de vida teve grande frequência de caméfitos (29.7%), nanofanerófitos (17.1%) e geófitos (15.3%). Entre as espécies de ocorrência restrita, geófitos (32.6%), caméfitos (25.6%) e nanofanerófitos (20.9%) como foram as mais frequentes. Testes de 2 indicaram diferenças significativas nas frequências de famílias e de formas de vida entre os ambientes. Nossos resultados indicaram uma associação positiva entre a cobertura de bromélias e a riqueza e a abundância de plântulas de espécies arbóreas. Aparentemente, maiores coberturas por B. balansae fornecem proteção contra a herbivoria, indistintamente, e amenização das severas condições ambientais, favorecendo o estabelecimento de espécies pioneiras da floresta. Os testes multivariados indicaram efeito significativo do fator sítio sobre a diversidade beta, em todas as classes de tamanho, e do fator gradiente somente para duas classes de tamanho analisadas. A vegetação rupestre apresentou uma flora distinta, florística e funcionalmente, com uma grande similaridade a outras formações rupestres neotropicais, mas com expressivo número de espécies arbóreas. Plântulas destas espécies, aparentemente, beneficiam-se da presença de B. balansae, o que pode levar à redução gradual da vegetação rupestre neste mosaico floresta/afloramentos. Entender como as comunidades arbóreas têm sua distribuição determinada por limitações de dispersão e de nicho pode levar a uma melhor compreensão dos processos que geram e mantêm a diversidade. / Rocky outcrops are centers of taxonomic and functional diversity with extreme environmental conditions. They also are sites for colonization by forestry species, usually related to facilitation mechanisms. The species distribution set by niche and neutrals mechanisms results in the variation of composition among communities (beta diversity). The goals of this work are evaluate if rocky outcrops are taxonomically and functionally different from the surrounding forest, study the possible positive interactions among species related to the advance of the forest over the rupestrian vegetation and determine how the composition of arboreal species in the interface forest/rocky outcrop is limited by neutral, niche or both dynamics along the ontogeny. We studied the vegetation of basaltic rocky outcrops at the Turvo State Park, south Brazil. The flora from rupestrian communities was analyzed at field during two years. Species were classified into life forms based on literature and field observations. To evaluate the patterns of arboreal beta diversity and the occurrence of facilitation between plants we sampled arboreal species in three forest/rocky outcrop ecotones. In each rocky outcrop we established 60 plots of 1.5 x 1.5 m along the forest-rocky outcrop ecotone. We described each plot in terms of Bromelia balansae Mez cover and the composition of tree species, separated into four size classes, representing different ontogenetic stages of individuals. The taxonomic and functional differences between the rupestrian vegetation and regional flora were analyzed with the chi-square test and the effects of different factors were analyzed with randomization tests uni-and multivariate. We registered 111 species, 43 of which restricted to the rocky outcrops, distributed in 54 botanic families. The life form spectrum showed a high proportion of chamaephytes (29.7%), nanophanerophytes (17.1%) and geophytes (15.3%). The flora restricted to rocky outcrops showed more frequently geophytes (32.6%), chamaephytes (25.6%) and nanophanerophytes (20.9%). Chi-square tests indicated significant differences in frequencies among environments, for both families and life forms. Our results showed that the coverage of bromeliads is positively correlated to the richness and abundance of seedlings of arboreal species. It seems that higher cover by B. balansae gives protection against herbivory indistinctively, and also reduces the effects of severe environmental conditions, allowing the settlement of forest pioneers species. Multivariate tests showed significant effect of the site factor on the beta diversity in all classes of size and significant effect of the gradient factor only on two size classes evaluated. The rupestrian vegetation is distinct floristically and functionally, showing high floristic similarity with other neotropical rupestrian formations, but has high number of arboreal species. Seedlings of arboreal species seem to have benefit from the presence of B. balansae, which can cause a gradual reduction of the rupestrian vegetation in the forest and rocky outcrop mosaic. Understanding how the distribution of arboreal communities is determined by limitations of dispersal and niche can improve the comprehension of the dynamics that generate and maintain diversity.

Marine Inseln beherbergen einen großen Teil der biologischen Vielfalt unseres Planeten und weisen gleichzeitig einen hohen Anteil endemischer Arten auf. Inselbiota sind allerdings zudem besonders anfällig für anthropogene Einflüsse wie den globalen Klimawandel, Habitatverlust und invasive Arten. Für ihren Erhalt ist es daher wichtig, die ökologischen Prozesse auf Inseln detailliert zu verstehen. Aufgrund ihrer definierten Größe und isolierten Lage eignen sich Inseln als Modellsysteme in der ökologischen und evolutionären Forschung. Der Großteil der bisherigen Inselstudien hat sich allerdings mit kleinräumigen Mustern befasst, so dass standardisierte globale Daten zu den biogeographischen Eigenschaften und eine makroökologische Synthese ihrer Biota bislang fehlen. In dieser Arbeit stelle ich eine physische und bioklimatische Charakterisierung der Inseln der Welt vor und behandle die Frage, wie abiotische Inseleigenschaften die Diversität von Inselfloren beeinflussen. Ich bearbeite zwei Hauptaspekte dieser Fragestellung: Zuerst konzentriere ich mich auf historische und heutige Klimabedingungen und physische Inseleigenschaften als Triebfedern von Pflanzendiversitätsmustern auf Inseln. Hierbei setze ich einen Schwerpunkt auf die räumliche Anordnung von Inseln und Struktur von Archipelen. Als Zweites behandle ich taxon-spezifische Unterschiede in der Antwort von Diversitätsmustern auf abiotische Faktoren. Hierzu stelle ich eine globale Datenbank mit historischen und heutigen Klimabedingungen und physischen Eigenschaften, wie Fläche, Isolation und Geologie, von 17883 Inseln größer als 1 km² vor. Mit Hilfe von Ordinations- und Klassifikationsverfahren charakterisiere und klassifiziere ich die Inseln in einem multidimensionalen Umweltraum. Außerdem entwickele ich einen Satz von ökologisch relevanten Maßen zur Beschreibung von Isolation von Inseln und ihrer räumlichen Anordnung in Archipelen, darunter Maße zu Trittstein-Inseln, Wind- und Meeresströmungen, klimatischer Ähnlichkeit, Distanzen zwischen Inseln und umgebender Landfläche. Diese Maße berücksichtigen verschiedene Aspekte von Isolation, welche Immigration, Artbildung und Aussterben auf Inseln sowie Austausch zwischen Inseln beeinflussen. Um abiotische Bedingungen mit biotischen Eigenschaften von Inselfloren in Verbindung zu bringen, nutze ich eine für diese Arbeit erstellte Datenbank aus 1295 Insel-Artenlisten, die insgesamt ca. 45000 heimische Gefäßpflanzenarten umfassen. Dies ist der umfassendste und erste globale Datensatz für Pflanzen auf Inseln, der Artidentitäten anstatt lediglich Artenzahlen beinhaltet. Die globale Insel-Charakterisierung bestätigt quantitativ, dass sich Inseln in bioklimatischen und physischen Eigenschaften vom Festland unterscheiden. Inseln sind im Durchschnitt signifikant kühler, feuchter und weniger saisonal geprägt als das Festland. Die weiteren Ergebnisse zeigen, dass eine sorgfältige Beschreibung der räumlich-physischen Eigenschaften von Inseln und Archipelen nötig ist, um die Diversitätsmuster ihrer Biota zu verstehen. Isolation ist nach Inselfläche der zweitwichtigste Einflussfaktor für den Gefäßpflanzenartenreichtum auf Inseln. Von den verglichenen Isolationsmaßen eignet sich der Anteil an umgebender Landfläche am besten zur Erklärung der Artenzahlen. Außerdem erhöht sich durch die Berücksichtigung von Trittsteininseln, großen Inseln als Quell-Landflächen und klimatischer Ähnlichkeit der Quell-Landflächen die Vorhersagekraft der Modelle. Isolation spielt eine geringere Rolle auf großen Inseln, wo in situ Diversifizierung den negativen Effekt von Isolation auf Immigration ausgleicht. Die räumliche Struktur innerhalb von Archipelen ist von besonderer Bedeutung für β-Diversität, d.h. für den Unterschied in der Artenzusammensetzung der Inseln. Außerdem beeinflusst sie indirekt, durch den Effekt auf die β-Diversität, auch die γ-Diversität, d.h. die Diversität des gesamten Archipels. Die Ergebnisse heben die enorme Bedeutung der relativen räumlichen Position von Inseln zueinander für Diversitätsmuster auf Inseln hervor und zeigen die Notwendigkeit für Inselforschung und Naturschutz, Inseln im Kontext ihres Archipels zu betrachten. Die Ergebnisse für Farne auf südostasiatischen Inseln zeigen, dass die Bedeutung von physischen Inseleigenschaften für Diversität kontinuierlich mit der Größe der betrachteten Untersuchungsfläche von der Insel- bis zur Plotebene abnimmt, wohingegen der Einfluss von lokalen Umweltbedingungen zunimmt. Lokale Artgemeinschaften sind häufig gesättigt, wodurch die Anzahl an Arten, die aus dem regionalen Artenbestand einwandern können, limitiert wird. Um Vorhersagen über lokalen Artenreichtum zu machen, ist es daher wichtig, die Skalenabhängigkeit der Effekte des regionalen Artenbestandes zu berücksichtigen. Großgruppen von Pflanzen unterscheiden sich in ihrer Ausbreitungsfähigkeit, ihrem Genfluss, Artbildungsraten und Anpassungen an das Klima. Dementsprechend zeigen die vergleichenden Analysen zwischen taxonomischen Pflanzengruppen deutliche Unterschiede in der Reaktion von Artenreichtum und phylogenetischen Diversitätsmustern auf abiotische Faktoren. Die Arten-Fläche-Beziehung, d.h. die Zunahme von Artendiversität mit zunehmender Fläche, variiert zwischen den Pflanzengruppen. Die Steigung der Arten-Fläche-Beziehung ist für Spermatophyten größer als für Pteridophyten und Bryophyten, wohingegen der y-Achsenabschnitt kleiner ist. Unter der Annahme, dass Merkmale und klimatische Anpassungen innerhalb von taxonomischen Gruppen phylogenetisch konserviert sind, führen die Filterwirkung von Ausbreitungsbarrieren und Umwelteigenschaften sowie in situ Artbildung zu Gemeinschaften eng verwandter Arten (phylogenetic clustering). Die Ergebnisse zeigen, dass physische und bioklimatische Inseleigenschaften, die mit der Filterwirkung und Artbildung in Verbindung stehen, die phylogenetische Struktur von Inselgemeinschaften beeinflussen. Die Stärke und Richtung der Zusammenhänge variieren zwischen taxonomischen Gruppen. Abiotische Faktoren erklären mehr Variation in phylogenetischer Diversität für alle Angiospermen und Palmen als für Farne, was auf Grund höherer Ausbreitungsfähigkeit und größerer Verbreitungsgebiete von Farnen den Erwartungen entspricht. Die abiotische Charakterisierung und Klassifizierung der weltweiten Inseln und die zugehörigen Daten ermöglichen eine integrativere Berücksichtigung von Inseln in der makroökologischen Forschung. In dieser Arbeit präsentiere ich die ersten Vorhersagen globaler Pflanzenartenvielfalt auf Inseln und die ersten Analysen zu unterschiedlichen Diversitätskomponenten (α, β, γ und phylogenetische Diversität) von Inselsystemen und ihren abiotischen Einflussfaktoren auf globalem Maßstab. Ich zeige, dass Zusammenhänge zwischen Umweltfaktoren und Artenzahl sowie phylogenetischen Eigenschaften von Inselgemeinschaften zwischen unterschiedlichen taxonomischen Gruppen in Abhängigkeit ihrer vorwiegenden Ausbreitungs- und Artbildungseigenschaften variieren können. Dies ist eine neue Sichtweise in der makroökologischen Inselforschung, die Rückschlüsse auf die Mechanismen hinter Diversitätsmustern von Pflanzen auf Inseln erlaubt. Ein detailliertes Verständnis davon, wie Diversität unterschiedlicher Pflanzengruppen durch Immigration und Diversifizierung auf Inseln entsteht, dürfte auch das Verständnis globaler Diversitätsmuster im Allgemeinen verbessern.