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.

Les activités humaines, de loisir et professionnelles, vont en s’intensifiant sur les zones rocheuses intertidales. Le piétinement associé à ces activités, le remaniement des blocs pour rechercher des espèces consommables, ne sont pas sans incidence sur les biocénoses des milieux rocheux. Les conséquences de ces activités sur la structure et la dynamique des roches du médiolittoral et des champs de blocs de bas d’estran en particulier, nécessitent d'être caractérisées. A travers l’identification des espèces et descripteurs qui répondent le plus aux perturbations "piétinement des platiers rocheux" et "retournement des blocs par les pêcheurs à pied", des indicateurs écologiques simplifiés peuvent être dégagés et faciliter, à terme, le diagnostic de l’état de conservation de ces habitats. Les processus écologiques de dégradation et de déstructuration des biocénoses sous l'effet du piétinement passent par des interactions entre espèces très complexes. Les résultats d’expérimentations in situ soulignent des phénomènes de seuils très marqués, liés en particulier à la protection jouée par les algues brunes structurantes sur les autres espèces. L'étude du retournement des blocs par les pêcheurs à pied met en évidence une phase de mortalité presque totale des organismes fixés ou peu mobiles des blocs et une faible résilience de l’habitat. Le choix de stations de référence appropriées et l’utilisation des espèces et descripteurs les plus sensibles au retournement des blocs ont permis la création de deux indicateurs de perturbation. Déclinés à deux échelles spatiales différentes, ils sont applicables en routine par les scientifiques et les gestionnaires d’aires marines protégées. / Recreational and professional activities are intensifying on the rocky intertidal. The overturning of boulders and the trampling associated with human activities, have consequences on the structure and dynamic of rocky intertidal communities. Through the identification of species and descriptors that respond more to the disturbances “trampling of mediolittoral rock platforms” and “the overturning of low eulittoral boulders by hand-fishermen”, SMART indicators (Simple, Measurable, Achievable, Realistic, and Time limited indicator) can be created for the assessment of the conservation status of these habitats. The effects of trampling were tested with control approach. Experiments in situ underlined very complex interactions between species during the ecological processes of degradation of biocenoses. High thresholds of disturbance were also observed. They were particularly related to the protection played by structuring brown algae on other species. The study of boulder’s overturning highlighted a step of almost total mortality of fixed or low mobile organisms that live on upper or lower surfaces of boulders. Results also showed a low resilience of the habitat. The use of appropriate reference situations and of the most sensitive species to the boulder’s overturning, led to the creation of two ecological indicators. Declined to two different spatial scales, they may be applied routinely by scientists and managers of marine protected areas.

This study is based at Rocky Canyon Quarry (RCQ), a 200-acre granite aggregate open-pit quarry with chaparral-dominated plant communities located in San Luis Obispo County, CA. At RCQ, the Surface Mining and Reclamation Act (SMARA) of 1975 was interpreted as restoring the landscape to native plant communities. Native plant community restoration projects have occurred there since 1993 through cooperation with California Polytechnic State University Biology Department in San Luis Obispo, CA. I evaluated past restoration at RCQ and researched new techniques to improve chaparral restoration based on the natural processes of fire. Chaparral is an important fire-dominated plant community within the California Floristic Province, which covers about seven percent of California. Typically during a fire, heat immediately acts on Adenostoma fasciculatum (Chamise) seeds/m2 in the soil seed bank. Smoke also reaches seeds on and near the soil surface. Chemical effects of fire, such as smoke and charcoal, are deposited on the soil surface and leach into the seed bank after fall rains. In nature, this results in enhanced germination of the seeds and the beginning of chaparral post-fire succession. Fire effects, both heat and chemical, have been supported to increase seed germination in numerous laboratory and field studies. I sought to utilize natural fire cues, such as heat, charate, and liquid smoke, to develop successful and efficient restoration prescriptions. The most successful restoration technique developed utilized Wright’s Liquid Smoke and heat to increase seed germination of Adenostoma fasciculatum (Chamise), Ceanothus cuneatus (California lilac), and Salvia mellifera (Black Sage) significantly. A new restoration prescription for RCQ based on literature reviews and the above mentioned research is presented.

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Most climate change predictions focus on the response of individual species to changing local conditions and ignore species interactions, largely due to the lack of a sound theoretical foundation for how interactions are expected to change with climate and how to incorporate them into climate change models. Much of the variability in species interaction strengths may be governed by fundamental constraints on physiological rates, possibly providing a framework for including species interactions into climate change models. Metabolic rates, ingestion rates and many other physiological rates are relatively predictable from body size and body temperature due to constraints imposed by the physical and chemical laws that govern fluid dynamics and the kinetics of biochemical reaction times. My dissertation assesses the usefulness of this framework by exploring the community-level consequences of physiological constraints. In Chapter 2, I incorporated temperature and body size scaling into the biological rate parameters of a series of realistically structured trophic network models. The relative magnitude of the temperature scaling parameters affecting consumer energetic costs (metabolic rates) and energetic gains (ingestion rates) determined how consumer energetic efficiency changed with temperature. I systematically changed consumer energetic efficiency and examined the sensitivity of network stability and species persistence to various temperatures. I found that a species' probability of extinction depended primarily on the effects of organismal physiology (body size and energetic efficiency with respect to temperature) and secondarily on the effects of local food web structure (trophic level and consumer generality). This suggests that physiology is highly influential on the structure and dynamics of ecological communities. If consumer energetic efficiency declined as temperature increased, that is, species did best at lower temperatures, then the simulated networks had greater stability at lower temperatures. The opposite scenario resulted in greater stability at higher temperatures. Thus, much of the community-level response depends on what species energetic efficiencies at the organismal-level really are, which formed the research question for Chapter 3: How does consumer energetic efficiency change with temperature? Existing evidence is scarce but suggestive of decreasing consumer energetic efficiency with increasing temperature. I tested this hypothesis on seven rocky intertidal invertebrate species by measuring the relative temperature scaling of their metabolic and ingestion rates as well as consumer interaction strength under lab conditions. Energetic efficiencies of these rocky intertidal invertebrates declined and species interaction strengths tended to increase with temperature. Thus, in the rocky intertidal, the mechanistic effect of temperature would be to lower community stability at higher temperatures. Chapter 4 tests if the mechanistic effects of temperature on ingestion rates and species interaction strengths seen in the lab are apparent under field conditions. Bruce Menge and I related bio-mimetic estimates of body temperatures to estimates of per capita mussel ingestion rates and species interaction strengths by the ochre sea star Pisaster ochraceus, a keystone predator of the rocky intertidal. We found a strong, positive effect of body temperature on both per capita ingestion rates and interaction strengths. However, the effects of season and the unique way in which P. ochraceus regulates body temperatures were also apparent, leaving room for adaptation and acclimation to partially compensate for the mechanistic constraint of body temperature. Community structure of the rocky intertidal is associated with environmental forcing due to upwelling, which delivers cold, nutrient rich water to the nearshore environment. As upwelling is driven by large-scale atmospheric pressure gradients, climate change has the potential to affect a wide range of significant ecological processes through changes in water temperature. In Chapter 5, my coauthors and I identified long-term trends in the phenology of upwelling events that are consistent with climate change predictions: upwelling events are becoming stronger and longer. As expected, longer upwelling events were related to lower average water temperatures in the rocky intertidal. Furthermore, recruitment rates of barnacles and mussels were associated with the phenology of upwelling events. Thus climate change is altering the mode and the tempo of environmental forcing in nearshore ecosystems, with ramifications for community structure and function. Ongoing, long-term changes in environmental forcing in rocky intertidal ecosystems provide an opportunity to understand how temperature shapes community structure and the ramifications of climate change. My dissertation research demonstrates that the effect of temperature on organismal performance is an important force structuring ecological communities and has potential as a tractable framework for predicting the community level effects of climate change. / Graduation date: 2013 / Access restricted to the OSU Community, at author's request, from Nov. 20, 2012 - Nov. 20, 2014

Subalpine fir (Abies lasiocarpa [Hooker] Nuttall) and white spruce (Picea glauca [Moench] Voss) trees were sampled in an old growth forest in the northern Canadian Rocky Mountains. Dendroclimatological methods were used to analyse the relationship between annual radial-growth and climatic variability. The white spruce ring-width chronology showed stronger sensitivity to climatic variability than the subalpine fir chronology. Both chronologies were positively correlated with growing season mean and minimum temperature. Additionally, the white spruce chronology was correlated with summer maximum temperature, late spring minimum temperature, and diurnal temperature range during the growing season. The subalpine fir ring-width chronology was also correlated with maximum and minimum temperature and diurnal temperature range during the during the previous winter and with the Pacific Decadal Oscillation during each month from December to June. Analysis of the climate-growth responses of individual trees revealed a higher level of intraspecies variability in subalpine fir than in white spruce. The white spruce chronology was selected for use in creating a proxy climate record based on its greater length and stronger sensitivity to climatic variability. Dendroclimatological methods were used to create a regional proxy record of June-July mean temperature extending back to 1772. This reconstruction exhibits a shared pattern of low-frequency variability with other dendroclimatic reconstructions from western Canada and shows no evidence of the recent reduction in sensitivity to climatic variability that is apparent in many other northern spruce chronologies. This study represents the first detailed dendroclimatic analysis undertaken in northern interior British Columbia. This work has elucidated the complex interactions between climate and the radial growth of alpine conifers in the northern Canadian Rocky Mountains. The climate reconstruction presented here fills in one of the remaining spatial gaps in the coverage of annually resolved climate reconstructions in western North America.

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