Crayfish throughout North America, Europe, and Asia host assemblages of obligate ectosymbiotic annelid worms called branchiobdellidans. The work presented here is a detailed experimental and observational study of the ecological interactions between crayfish and their worms. In a comprehensive literature review, I show that branchiobdellidans have complex and context-dependent effects on their hosts, serving as both beneficial cleaners and tissue-consuming parasites. Using a field survey and laboratory experiments, I provide novel evidence for age-specific resistance as an adaptation to maximize life-long benefits of a mutualism. Specifically, I show that Cambarus crayfish display a consistent ontogenetic shift in resistance to the colonization of branchiobdellidans and this shift likely reflects underlying changes in the costs and benefits of symbiosis. I then show that this change in host resistance creates predictable patterns of symbiont diversity and composition throughout host ontogeny. Host resistance limits within-host symbiont communities to a few weakly interacting species, whereas relaxed resistance leads to more diverse symbiont communities that have strong interactions among symbiont taxa. Thus, host resistance has direct effects on within-host symbiont community structure by selectively filtering colonizing species, and indirect effects by moderating the strength of interactions among symbionts. Lastly, in a detailed study of the worm Cambarincola ingens, I depict a symbiont dispersal strategy that yields highly predictable transmission dynamics during pairwise host-host encounters and shows that variation in transmission dynamics can be explained by the fitness outcomes for dispersing symbionts. Field observations revealed that worm reproduction is contingent on host size and intraspecific competition for preferred microhabitats. Using a predictive model that assumes transmission of symbionts only when current conditions yield fitness below a minimum threshold, I was able to predict individual transmission events much more accurately than a comparable null model that assumed a fixed probability of transmission. My work provides empirical support for the emerging trend among researchers that advocates the adaptation of general ecological frameworks to understand symbiont population structure and diversity, but my work also emphasizes the value of detailed natural history studies to uncover system-specific ecological and co-evolutionary processes such as partner control mechanisms, symbiont microhabitat selections, and symbiont dispersal strategies. / Ph. D.

Global change is altering the community composition, variability, and behavior of organisms in a diverse suite of ecosystems. Because of climate change and eutrophication, freshwater lakes and reservoirs are experiencing an increase in low dissolved oxygen concentrations (hypoxia) in their bottom waters (hypolimnion), which can disrupt ecological communities. Zooplankton, important aquatic organisms for regulating water quality and food webs, are one group of organisms affected by hypoxia since zooplankton need oxygen to respire. My research shows that hypoxia may disrupt zooplankton behavior and increase the variability of zooplankton communities. Zooplankton ubiquitously exhibit diel vertical migration, where the majority of the population resides in the hypolimnion during the daytime to escape predation from fish and damage from ultraviolet radiation. At night, many zooplankton ascend to the surface waters to feed on phytoplankton, when there is decreased risk of predation and radiation. My results from intensive 24-hour sampling campaigns suggest that hypolimnetic hypoxia may alter zooplankton migration, biomass, and behavior, which may in turn exacerbate water quality degradation due to the critical role zooplankton play in freshwater ecosystems. In addition, field surveys in four reservoirs over three years revealed that hypoxia may increase the variability of zooplankton communities compared to oxic conditions. Consequently, as lakes and reservoirs experience increased extent and duration of hypoxia in the future, it is critical to understand how more variable zooplankton communities alter freshwater ecosystem functioning. / Ph. D. / The globalization and extensive use of habitats by humans has created stress on other organisms in many ecosystems; the interactions, migration patterns, and composition of species have been altered globally because of humans. In freshwater lakes and reservoirs, warmer water temperatures and increased nutrient pollution have resulted in decreased oxygen concentrations in many waterbodies. Decreased oxygen concentrations can affect the organisms living in these environments that need oxygen to survive, which can disrupt the water quality and function of the ecosystem. Zooplankton, or animal plankton or animal drifters, are one group of organisms affected by low oxygen conditions. Zooplankton play an important role in aquatic ecosystems. Zooplankton eat phytoplankton (at the base of the food chain) and are consumed by fish; therefore, any changes to zooplankton populations from low oxygen concentrations can affect water quality and fisheries. My research shows that low oxygen concentrations affect the migration patterns of zooplankton. Zooplankton generally migrate vertically in the water column of a lake and reservoir over each 24-hour period to optimize their proximity to phytoplankton (their food source), while minimizing their chance of being eaten by fish. I found that low oxygen concentrations disrupt this cycle; zooplankton largely do not vertically migrate under low oxygen concentrations. My results from intensive 24-hour sampling campaigns and sampling over multiple summer periods suggest that low oxygen concentrations disrupt zooplankton migration and behavior, which may exacerbate water quality issues in lakes and reservoirs.

<p>Climate change has dramatically altered the ecological landscape of the eastern U.S., leading to shifts in phenological events and redistribution of tree species. However, shifts in phenology and species distributions have implications for the productivity of different populations and <a></a>the communities these species are a part of. Here, I utilized two studies to quantify the effects of climate change on forests of the eastern U.S. First, I used phenology observations at a common garden of 28 populations of northern red oak (<i>Quercus rubra</i>) across seven years to assess shifts in phenology in response to warming, identify population differences in sensitivity to warming, and correlate sensitivity to the productivity of the populations. Second, I utilized data from the USDA Forest Service’s Forest Inventory and Analysis Program to identify forest communities of the eastern U.S., assess shifts in their species compositions and spatial distributions, and determine which climate-related variables are most associated with changes at the community level. In the first study, I found that populations were shifting their spring phenology in response to warming, with the greatest sensitivity in populations from warmer, wetter climates. However, these populations with higher sensitivity did not have the highest productivity; rather, populations closer to the common garden with intermediate levels of sensitivity had the highest productivity. In the second study, I found that there were 12 regional forest communities of the eastern U.S., which varied in the amount their species composition shifted over the last three decades. Additionally, all 12 communities shifted their spatial distributions, but their shifts were not correlated with the distance and direction that climate change predicted them to shift. Finally, areas with the highest changes across all 12 communities were associated with warmer, wetter, lower temperature-variable climates generally in the southeastern U.S. Taken together, these studies provide insight into the ways in which forests are responding to climate change and have implications for the management and sustainability of forests in a continuously changing global environment.</p>

Thesis (PhD)--Stellenbosch University, 2015. / ENGLISH ABSTRACT: Understanding how divergent selection results in the evolution of reproductive isolation (i.e. speciation) is an important goal in evolutionary biology. Populations of herbivorous insects using different host plant species can experience divergent selection from multiple selective pressures which can rapidly lead to speciation. Restio leafhoppers are a group of herbivorous insect species occurring within the Cape Floristic Region (CFR) of South Africa. They are specialised on different plant species in the Restionaceae family. Throughout my thesis I investigated how bottom-up (i.e. plant chemistry/morphology of host plant species) and top-down (i.e. predation and competition) factors drive specialisation and divergence in restio leafhoppers. I also investigated interspecific competition as an important determinant of restio leafhopper community structure. In chapter 2 I quantified host specificity of restio leafhopper species within a local community for 24 months. I found that restio leafhopper species are highly host specific and potentially synchronised with the growth phases of their host plants. In chapter 3 I used a network metric, modularity, to determine whether host plant partitioning in a restio leafhopper community is non-random (i.e. driven by a deterministic process). This metric allows the identification of the components underlying host plant partitioning (modules). I then performed experiments to show that modules, and therefore host plant partitioning, can mostly be explained by preference and performance relationships (i.e. bottom-up factors). In chapter 4 I used null models to test whether niche partitioning in restio leafhopper communities is a general pattern across the landscape. I found non-random niche partitioning, which results from strong host specificity, in all investigated restio leafhopper communities. In addition, I performed binary host choice experiments in the presence and absence of interspecific competition, but found no evidence that interspecific competition narrows host preferences. These findings suggest that host specificity, the cause of niche partitioning, is likely shaped over evolutionary time. Sampling multiple interaction networks across the CFR, in chapter 5, I tested whether restio leafhopper populations are more host specific in species rich communities and regions in the CFR than in species poor communities and regions. I found no positive relationship between restio leafhopper species richness and host specificity at any scale. These findings suggest that specialisation is not driven by interspecific competition. In chapter 6 I investigated host shifts in Cephalelus uncinatus. C. uncinatus has a broader distribution than any single restio species that it can use; suggesting that host plant related divergence may result from geographic range expansion. I found that allopatric and parapatric populations, but not sympatric individuals, using different host plants have divergent host preferences. I also found evidence for morphological divergence in traits related to predator avoidance in population pairs that exhibit divergent host preferences. My findings emphasise the importance of both bottom-up and top-down factors, with the exception of interspecific competition, as determinants of specialisation and divergence in restio leafhoppers. I find no evidence that interspecific competition is an important force structuring restio leafhopper communities. Instead, strongly niche partitioned community structure appears to emerge from the speciation process. / AFRIKAANSE OPSOMMING: Die wyse waarop uiteenlopende seleksie lei tot die evolusie van seksuele isolasie (n.l. spesiasie) is ‘n belangrike vraag in evolutionêre biologie. Plantetende insekpopulasieses wat verskillende gasheerplante gebruik kan onder uinteenlopende veelvoudige seleksie wees en vinnig spesiasie ondergaan. Restio-blaarspringers is ‘n groep plantetende insekspesies wat gespesialiseerd is op verskillende plantspesies in die restio familie. In my tesis ondersoek ek die onder-op (n.l. plantchemie en morfologie) en bo-af seleksiekragte (n.l. predasie en kompetisie) wat lei tot gasheerspesialisasie en -spesiasie in restio-blaarspringers. Ek ondersoek ook die belangrikheid van tussen-spesieskompetisie in gemeenskapsorganisasie. In hoofstuk 2 het ek gasheerspesialisasie gekwantifiseer in ‘n klein restio-blaarspringergemeenskap oor 24 maande. Ek het gevind dat restio-blaarspringers hoogs gasheerspesifiek is en moontlik met die groeifase van hul hoofgasheerplante gesinchroniseerd is. Ek het in hoofstuk 3 ‘n netwerkmetriek, modulariteit, gebruik om te bepaal of restio-blaarspringers se gasheerverdeling nie-stokasties is (n.l. deur deterministiese prosesse veroorsaak is). Hierdie metriek laat ‘n mens toe om die komponente van gasheerverdeling (modules) te identifiseer. Deur middel van eksperimente het ek bepaal dat modules, en dus gasheerverdeling, deur gasheervoorkeur en prestasie (onder-op prossesse) verduidelik kan word. In hoofstuk 4 het ek ondersoek of gasheerverdeling algemeen is. Ek het deur middel van nulmodelle gewys dat gasheerverdeling algemeen is en veroorsaak is deur sterk gasheerspesifiekheid. Ek het ook voorkeureksperimente uitgevoer in die teenwoordigheid en afwesigheid van tussen-spesies kompetisie. Hier het ek geen teken gevind dat huidige tussen-spesies kompetisie gasheervoorkeur beïnvloed nie. My bevindinge in hierdie hoofstuk stel dus voor dat spesialisasie, die oorsaak van gasheerverdeling, oor evolutionêre tyd gevorm word. In hoofstuk 5 het ek ondersoek of populasies van restio-blaarspringers meer gasheerspesifiek is in restio-blaarspringerspesies ryke gemeenskappe en streke as populasies in spesies-arm gemeenskappe en streke. Ek het geen positiewe korrelasie tussen spesiesrykheid en gasheerspesialisasie gevind nie. Dit dui daarop aan dat gasheerspesialisasie, en dus gasheerverdeling, nie deur tussen-spesies kompetisie veroorsaak word nie. In hoofstuk 6 het ek gasheerplantgekoppelde divergensie ondersoek in Cephalelus uncinatus. C. uncinatus se verspreiding is breër as enige restio-spesies wat dit kan gebruik. Dit stel voor dat verspreidingvergroting gasheerverskuiwing mag veroorsaak. Ek het gevind dat populasies wat verskillende plante in allopatrie en parapatrie gebruik uiteenlopende gasheerkeuses maak, maar insekte wat verskillende plante in sympatrie gebruik wys nie ontwrigtende gasheerkeuses nie. Die populasies wat uiteenlopende gasheerkeuses getoon het, het ook verskillende morfologiese teen-predasie eienskappe getoon. Dit dui daarop aan dat predasie belangrik mag wees vir spesiasie in restio-blaarspringers. Die bevindinge van my tesis dui daarop aan dat beide onder-op en bo-af seleksie belangrik is vir gasheer-spesialisasie in divergensie. Nietemin, tussen-spesies kompetisie is nie ‘n belangrike bron vir spesialisasie, divergensie of gemeenskapsorganisasie nie. Gemeenskapsorganisasie is klaarblyklik slegs ‘n gevolg van die spesiasie proses.

The trait based approach has been proposed as a way to reconcile community ecology. Despite recent advances in trait based ecology, such as the development of global trait databases and standardized methodology for trait collections, it remains unclear to what degree traits vary across individuals, species, and communities. In addition, the drivers of trait variation may shed light on the underlying processes that maintain species diversity and community assembly at local to continental scales yet these have been poorly studied. In this study, I examine both the magnitude of trait variation as well as the patterns of trait variation at local to continental scales in order to understand the drivers of diversity patterns across environmental gradients. First, I quantified the magnitude of trait variation at local scales in a dry tropical forest and determined that intraspecific variation is not negligible and can be quite large for compound-leaved species. However, I showed that the sample sizes necessary for quantifying trait variation are tractable and should encourage the adoption of trait variation in trait based ecology. Second, I tested whether climatic variables are predominantly responsible for observed trait variation across dry tropical forests in the Americas. I showed that climatic variability, specifically variability in precipitation, explained a large degree of observed trait variation across dry tropical forests and may provide a unique approach for classifying dry tropical forests based on their inherent degree of climatic seasonality. Third, I quantified patterns of trait variation at continental scales across elevational gradients at high to low latitudes. I showed that climatic variables largely drive patterns of trait variation at high latitudes while biotic factors largely drive patterns of trait variation at low, tropical latitudes. This finding has implications for understanding large-scale patterns of species diversity across elevational and latitudinal gradients. Finally, I apply trait variation to life history theory by quantifying variation in two life history traits (growth and reproduction) in a tropical tree species using a legacy dataset. I showed that variation in these two life history traits is due to both resource availability and allometric related effects on both traits. In sum, this study advances our understanding of the magnitude and underlying drivers of trait variation at local to continental scales.

Predicting the impacts of non-native species remains one of the greatest challenges to invasion ecologists. Because of their insularity, freshwater systems are particularly vulnerable to invasions, especially from non-native predators. The research in this thesis explores the role of abiotic and biotic factors in mediating the distribution and impact of Bythotrephes longimanus, a predatory cladoceran that has been introduced to freshwater systems in North America. Although the general impacts of this invasion have been documented, little is known about the factors that modulate them. Using a combination of field surveys and experiments, I tested several hypotheses concerning the influence of interactions with native species, as well as the role of heterogeneity in the light environment, in mediating the impact of Bythotrephes. Results demonstrated that biotic resistance by native macroinvertebrate predators does not play a limiting role in the establishment success of Bythotrephes. However, the within-lake distribution of Bythotrephes was influenced by these macroinvertebrates, suggesting that the native predator context matters when trying to understand the impacts of non-native predators. This was demonstrated with a mesocosm experiment in which the impact of Bythotrephes was constrained by the native Chaoborus larvae. In terms of the abiotic environment, in situ feeding experiments demonstrated that refuges from impact could exist for some prey taxa, as the outcome of predation by Bythotrephes was dependent on light availability and some prey taxa were more successful at evading predation under low light conditions. Finally, results show that adaptive behaviour by prey is also an important determinant of impact, as migrating Daphnia can escape predation effects by Bythotrephes. The combination of light-limited predation and a shallow distribution by Bythotrephes selects for prey that occupy relatively deeper positions during the day. As a whole, this research highlights the importance of complex interactions in mediating the impact of Bythotrephes and may help to explain some of the variation that has been documented among invaded lakes. A better understanding of these complex interactions can improve our ability to anticipate impacts as Bythotrephes continues to spread, as well as provide insight on some of the long-term effects following invasion. / Thesis (Ph.D, Biology) -- Queen's University, 2013-06-17 09:26:35.221

Food web structure is of underlying importance to ecological functions and processes. Whilst it is understood that a range of biotic and abiotic factors affect structure, relatively little is known of the role of biodiversity per se in structuring food webs. In this thesis I utilise novel multi-dimensional estimates of food web structure based on stable isotope ratios of nitrogen (δ15N) and carbon (δ13C) to quantify structural responses to changing community diversity. I additionally investigate methodological aspects of sample preparation and stable isotope quantifications of food chains. Using an arthropod prey-predator system, in chapter 2 I demonstrate that tissue selection and lipid extraction are important methodological procedures for deriving accurate δ15N and δ13C signatures. In chapter 3 I test the utility of δ15N to quantify food chain length, and δ13C to trace primary energy sources through to end consumers. Bayesian resampling of variance in sample means for plant and arthropod food chains produces robust isotopic estimates that match known food chain length well despite some error variance, and estimates of δ13C-range that trace trophic transfers. Chapter 4 represents a change in system from lab to field as I determine δ15N and δ13C signatures for plant and invertebrate species within three grassland communities representing a gradient of biodiversity. Quantifications of community bivariate isotopic space using isotopic metrics revealed that greater taxonomic richness increased both diversity of resource space exploited and overlap in resource space. These results therefore suggest that loss of diversity affected structure through altering relative patterns of niche partitioning in resource exploitation amongst community members. In chapter 5, I additionally find evidence that grassland management mediated changes in food web compartmental structure that were associated with differences in generalist invertebrate predator feeding habits. Taken together, these findings develop and demonstrate the utility of isotopic approaches to quantifying food web structure, and provide evidence of important mechanisms by which biodiversity affects food web structure. I conclude that the preservation of natural food web structure and trophic dynamics are further reasons for halting loss of biodiversity.

Species diversity varies dramatically across the surface of the Earth. A key step in the accumulation of species diversity is the ability of species to coexist in biological communities. Thus, identifying the mechanisms underlying community assembly is a major challenge for ecologists seeking to explain patterns in species diversity and composition. Recently some consensus has been reached on the set of processes that influence community assembly: speciation, demographic stochasticity, niche-based fitness trade-offs among species and dispersal. However, it is unclear how the importance of a particular process changes with spatial scale, which interactions exist among processes at large spatial scales and the extent to which niche-based resource partitioning among species explains differences in diversity among communities. Neotropical birds offer an ideal opportunity to address these uncertainties because of their high diversity and the existence of detailed information on their evolutionary history and ecology. In this thesis, I first use trait and phylogenetic metrics of community structure to show that both habitat filtering and interspecific competition shape community composition at the scale of individual bird territories (~1-2 ha). Second, I use simulations of community assembly to show that trait-based metrics of community structure outperform phylogenetic metrics for detecting niche-based community assembly, and that both sets of metrics often have low power when multiple processes influence community composition. Third, taking a trait-based, species-level approach, I show that both habitat filtering and interspecific competition influence species occurrence at regional scales (~75000 km²), and interact with dispersal ability so that their effect on species occurrence is increased for species with greater dispersal ability. Finally, using a combination of trait- and isotope-based methods to quantify resource partitioning, I show that species' niche widths do not change and niche overlap is reduced at high compared to low species richness. Taken together, these results suggest that both habitat filtering and interspecific competition (i.e. niche-based processes) influence community assembly from local to regional scales. However, at least at regional scales, the degree to which these processes are important for determining the occurrence of any given species depends on that species's dispersal ability. They also suggest, based on niche-based interspecific competition influencing community composition, that differences in species richness among communities are in part explained by differences among sites in the breadth of available niche space, not by increased ecological specialisation or niche overlap.

Aujourd'hui la perte et la transformation des habitats sont les principales menaces qui causent la diminution de la diversité biologique. A Madagascar, 90% des espèces végétales sont endémiques de l’île et la plupart des formations végétales sont actuellement fortement dégradées ou remplacées par des formations secondaires résultant des activités humaines. Le bois de tapia, dominé par l’espèce endémique de Madagascar Uapaca bojeri, est une formation végétale sclérophylle limitée aux Hautes Terres de l’île. Cette formation, adaptée et résistante au régime de feu naturel de la région, est aujourd'hui fragmentée, couvrant une surface équivalente à 132 255 ha au total. Afin d’augmenter la protection du bois de tapia, une nouvelle aire protégée (NAP) a été établie sur le Massif d’Ibity. L’état actuel de la végétation de bois de tapia est le résultat de l’interaction de facteurs comme le type de sol, le climat, les pratiques humaines traditionnelles et le feu. Bien que le feu soit un des phénomènes qui fasse partie de la dynamique de cette végétation, le régime de feu actuel est également une des causes de sa dégradation. L’objectif de cette thèse est donc d’étudier le rôle du feu sur le cycle démographique et certains processus importants pour l’installation et le recrutement. Ainsi, la germination, la phénologie et la résistance des plantules au feu ont été étudiées. Cette recherche montre que le problème actuel du bois de tapia est 1) la réduction de la floraison et de la fructification par les fréquences de feu élevées ; 2) la réduction du pourcentage de germination après l’exposition des graines à de hautes températures, et 3) la mortalité élevée des plantules après le passage d’un feu, en particulier quand la quantité de combustible est élevée. Ainsi l’installation et le recrutement des espèces ligneuses sont limités par le feu, qui a un effet négatif sur la régénération naturelle. La gestion du feu autour de l’aire protégée est ainsi nécessaire pour sa conservation / Currently loss and transformation of habitats are the main threats which cause the decrease of biological diversity. In Madagascar, 90% of plants species are endemic of the island and most of the plant formations types are at present strongly degraded or replaced by secondary formations resulting from human activities. Tapia woodland, dominated by the endemic tree Uapaca bojeri, is a sclerophyllous vegetation type limited in the Malagasy highlands. This vegetation type, adapted and resistant to the natural fire regime, is very fragmented today, covering a surface equivalent to 132 255ha on the island. In order to increase the protection of tapia woodland, a new protected area was established on Ibity massif. The current state of the woody vegetation is the result of the interaction of some factors as soil type, climate, human traditional practices and fire. Although fire is one of the phenomena that determine the dynamics of this vegetation, the current fire regime is also one of the main causes of degradation. The objective of this thesis is to study the role of fire on the demographic cycle and on some main processes for installation and recruitment. Thus, germination, plant phenology and seedlings resistance after burnt were studied. This research shows that the current problem of tapia woodland are 1) the reduction of flowering and fruiting by high fire frequencies; 2) the reduction of germination percent after seed exposure with high temperatures, and 3) high seedlings mortality after burnt, in particular when the quantity of fuel is raised. Installation and recruitment of woody species are limited by fire, which has a negative effect on natural regeneration. Fire management around the protected area is thus necessary for its conservation

This diploma thesis was focused on the community ecology of corticolous subaerial cyanobacteria and algae. Altogether 54 morphotypes were identified on the surface of the tree bark in the Nature Reserve Březina in České středohoří Mts., Czech Republic. Compared to publish data on tropical and subtropical habitats, high morphological diversity of these organisms in a temperate region was thus recognized. Coccoid green algae - most of them belonging to the class Trebouxio-phyceae - dominated. In a study focused on the algae community structure at small spatial scales (such as a single tree trunk), the temporal factors were found as the most important factor influencing the species composition. The height of a sample on trunks and type of bark roughness seemed to influence the species richness, yet the differences were not significant. The alpha-diversity and species composition of corticolous algal communities among individual trees were strongly influenced by host tree species. The pH of the bark surface also had significant impact on species composition. Generally, trees with lower pH of the bark surface (alder, spruce) had lower species richness, whereas trees with more alkaline bark surface (ash, beech, maple) had higher alpha-diversity values. Considering the generally efficient dispersal ability of...