<p>The seasonal response patterns of net photosynthesis and respiration (and nitrogenase activity in Nostoc) are described within a multivariate framework of temperature, moisture and light for the alpine and grassland crustaceous lichens Rhizocarpon superficiale and Caloplaca trachyphylla respectively and for the grassland surface cyanophyte Nostoc commune. These physiological responses are discussed in context of each species' boundary layer environment, with particular emphasis placed on interactions between environmental constancy and adoption of acclimation strategies.</p> <p>For R. superficiale the high frequency of thermal fluctuations experienced by hydrated thalli, sometimes on an hourly basis, precluded any strategy of seasonal acclimation. Instead, photosynthetic rates exhibited a broad temperature response, remaining near 1 mg CO₂ h⁻¹g⁻¹ from 1 up to 21°C, with no changes evident between seasonal responses. In marked contrast C. trachyphylla shows a distinct winter/summer pattern of photosynthetic acclimation. In winter months rates are optimal near 7°C, while in summer the temperature optima of net photosynthesis shift to 21°C. These changes correlate well with predictable seasonal microclimate events, particularly those associated with winter Chinook snowmelt periods. A third pattern of response was seen in N. commune, where no seasonal changes in response patterns were evident and both nitrogenase activity and net photosynthesis were maximal near 35°C. This response pattern allows maximum carbon gain and nitrogen fixation during spring and summer periods following precipitation events, while its more sheltered aspect reduces the importance of winter snowmelt periods.</p> / Doctor of Philosophy (PhD)

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Previous work has demonstrated that coflowering plant species (those that flower simultaneously in the same place) can potentially compete for pollination services. Competition for pollination among plant species can negatively impact their reproductive success. To minimise competition, plants can partition the activity of shared pollinators through releasing their floral resources at different times. Resource partitioning has been studied in several individual plant species, and some guilds of plants (e.g. African acacias), but little is known about temporal changes in resources at a community level. This thesis examines the spatiotemporal changes of floral resources at a community level and its implications for pollinator activity patterns. The temporal patterns of nectar and pollen provision of 70 plant species in two different plots were investigated at Mpala, in north central Kenya between 2004 and 2006. The communities studied showed that seasonal and daily microclimatic fluctuations significantly affect flowering patterns, times of flower opening, dehiscence and nectar production; and consequently the overall amount of pollen mass and nectar volume available at different times. I explored the effects of daily temporal changes in floral resource availability on pollinator activity patterns both in a guild of Malvaceae plant species, and at the community level through pollination webs. Detailed observations of daily patterns of resource provision and floral visitation in six Malvaceae plant species showed that plants can effectively avoid competition through attracting different pollinators, and via resource partition. Examination of daily changes in resource availability and the links portrayed in plant-visitor webs revealed that visitors move from one plant species to another, actively tracking changes in floral resource provision. These results suggest that in combination with physiological limitations imposed to the pollinators by temperature, bottom-up influences are a main force shaping daily pollinator activity patterns at a community level. Competition for pollination can only occur if plant species flower simultaneously and share pollinators within the same geographic area. Competition for pollination has been investigated in at least two African communities, but none of these studies have assessed the geographic spatial scale over which competition among coflowering species might happen. With the aim of measuring the geographic distance that pollinators visiting African acacias fly whilst foraging, I used molecular techniques to conduct paternity analysis as a proxy of how far pollen is carried away from particular trees within a population of Senegalia (Acacia) mellifera. The paternity analysis showed that pollinators move on average a maximum of 60 metres from the sampled trees, and that trees producing more flowers (resources) receive more visits, confirming that if sufficient resources are available pollinators can stay within relatively small geographic areas. Pollen movement only provides a partial measurement of the genetic neighbourhood of individual plants, because genes via seed dispersal can travel longer distances than those encompassed by the pollinators foraging areas. To investigate patterns of genetic variability in S. mellifera, I used microsatellites to conduct landscape genetic analyses including 25 adult populations and 9 seedling populations. Fourteen distinctive genetic clusters separated by four main geographic barriers were identified in the analyses. Significantly higher inbreeding was found in the seedling populations than in the adult populations. I discuss the possibility that this situation has been caused by anthropogenic exploitation and fragmentation of the adult stands.

Water is a critical resource for which plants compete in many terrestrial communities. In arid communities where water most limits plant growth, rainfall events occur in discrete, pulsed events. These pulses of water create highly variable soil moisture availabilities. Plant species respond differently to variation in soil water availabilities throughout a season and between years. How species vary in their responses to a range of water availabilities is thought to influence community and ecosystem properties. Many previously proposed hypotheses are not suitable to explain rapidly fluctuating resource availabilities or numerous input events throughout the growing season. This dissertation uses variation in water availability as a model resource to examine how species characteristics influence the process of exploitation competition within plant communities.Experiments were conducted to examine variation in growing season, exploitation competition between several pairs of co-occurring species in the Sonoran desert. Three separate studies evaluated several components of community dynamics thought to be influenced by exploitation competition. Spatial attributes of exploitation competition were assessed by measuring the performance of a deep-rooted species across the boundary of a natural expansion of a shallowly rooted species. Then, neighborhood composition was varied for species of similar growth-form to address the affects of species characteristics to shifts in abundances under field conditions. Lastly, species from the neighborhood composition study were placed under controlled, manipulated water availabilities to measure their fundamental operational conditions.Performances of plant species in all experiments were assessed using a combination of physiological and vegetative measurements, capturing the responses of the plants to both the dynamic growth conditions during the growing season, and integrated measures of plant performance post growth season. A shared preference was found for all species, where the performance of all species was greatest when water was most available in the soil profile. This work suggests the mechanism within a functional type by which plants coexist at various abundances is in part due to the variation in responses to temporal resource gradients. The variation in availability of resources and the species composition within the community should be considered in studies of competition between plant species.

Differences in reproductive strategies are a major factor influencing the patterns of genetic variability. Inbreeding and other non-recombining breeding systems can have profound effects on the efficacy of natural selection, which should be manifested in the patterns of genetic diversity within and between species. The impact of an organism’s breeding system can be investigated through a number of approaches. In this thesis, I use mathematical modelling, computer simulations, breeding schemes, quantitative life history measures, and molecular biological techniques to explore many of the consequences of breeding system evolution. Following a general introduction in Chapter 1, I explore the dynamics of transposable elements (TEs)—selfish mobile sequences of DNA that have deleterious effects upon their hosts. Sexual reproduction and recombination are important for constraining TE abundance, and in the absence of sex, an unchecked proliferation of TEs may cause a population to go extinct. In Chapter 2, I use a theoretical framework to analyze TE dynamics under asexual reproduction. Here, I show that while small populations are driven to extinction by element accumulation, large asexual populations can prevent this fate and be cured of vertically transmitted TEs. These results may help explain an "evolutionary scandal": the persistence of ancient asexual lineages, such as the bdelloid rotifers. In Chapter 3, I extend the computer simulations used in the previous chapter to explore the effects of reduced recombination on the distribution and abundance of TEs in sexual populations. I show that TEs become fixed as a result of Hill-Robertson effects in the form of Muller’s ratchet, but only in regions of extremely low recombination when excision is effectively absent and synergism between elements is weak. These results should help explain genomic patterns of TE distributions. In the remainder of the thesis, I turn to testing the genetic effects of androdioecy—the breeding system in which populations are comprised of separate male and hermaphrodite individuals—using the nematode Caenorhabditis elegans and related species. This unusual breeding system promotes high levels of inbreeding, yet males are maintained at appreciable frequencies. In Chapter 4, I measure lifehistory traits in the progeny of inbred versus outcrossed C. elegans and the related outcrossing species, C. remanei, to compare levels of inbreeding depression. I show that highly inbred C. remanei show dramatic reductions in brood size and relative fitness compared to outcrossed individuals, whereas pure strains of C. elegans performed better than crosses between strains, indicating outbreeding depression. The results are discussed in relation to the evolution of androdioecy and the effect of mating system on the level of inbreeding depression. Like C. elegans, C. briggsae reproduces by self-fertile hermaphrodites, and both species have similarly low levels of molecular diversity. But the global sampling of natural populations has been limited and geographically biased. In Chapter 5, I describe the first cultured isolates of C. elegans and C. briggsae from sub-Saharan Africa, characterize these samples for patterns of nucleotide polymorphism and vulva precursor cell lineage variation, and conduct a series of hybrid crosses in C. briggsae to test for genetic incompatibilities. With the new African isolates, I show distinct differences in levels of genetic and phenotypic diversity between the two species. Despite many similarities between C. elegans and C. briggsae, the results indicate that there may be more subtle, and previously unknown, differences in their natural histories. Finally, I return to the question of the impact of reduced recombination on TE dynamics in Chapter 6, by comparing population frequencies of TEs in natural populations of selfing and outcrossing Caenorhabditis species. I show that in the selfing species, C. elegans, transposons are less polymorphic and segregate at higher frequencies compared with the outcrossing species, C. remanei. Estimates of the intensity of selection based on the population frequencies of polymorphic elements suggest that transposons are selectively neutral in C. elegans, but subject to weak purifying selection in C. remanei. These results are consistent with a reduced efficacy of natural selection against transposable elements in selfing populations.

Mechanisms of tooth replacement distribute incongruently among extant gnathostomes (jawed vertebrates): a permanent tooth-generating dental lamina exists in chondrichthyans (cartilaginous fish) and tetrapods but not teleosts, whereas tooth shedding by basal hard tissue resorption occurs in tetrapods and teleosts but not chondrichthyans. Theories about the evolution of tooth development have been biased towards the chondrichthyan conveyor-belt replacement, since there has been no fossil evidence for the origin of osteichthyan (bony fish and tetrapods) tooth replacement until now. 3D virtual dissections with submicron-scale resolution, based on propagation phase contrast synchrotron microtomography (PPC-SRµCT), reveal the growth history of the dentitions of Andreolepis and Lophosteus, 423-Myr-old Silurian stem osteichthyans close to the common ancestor of tetrapods and teleosts. Their marginal jawbones and “tooth cushions” (possible homologues of coronoids) shed teeth by in situ cyclic basal resorption, the earliest examples of osteichthyan-style tooth replacement. The replacement cycles were site-autonomic, and occurred in broad irregular multi-row tooth fields, including at sites separated from the margin of the bone by intervening teeth, showing that the production of replacement teeth did not occur in a single deep dental lamina, but in pockets associated with each tooth, as in many teleosts. It suggests that the functionally and anatomically similar laminae of chondrichthyans and tetrapods are convergent. The marginal jaw bones of both genera carry an initial non-shedding dentition arranged in alternate transverse files, labial to the shedding tooth field, overgrown by later dermal ornament and probably not belonging to the oral domain, but bearing in vivo biting damage showing that they functioned as teeth. The most lingual of these odontodes have been resorbed apically and are overlain by shedding teeth. The first-generation teeth on the tooth cushions display basal resorption in Andreolepis, but semi-basal resorption in Lophosteus. The latter leaves a basal dentine ring from each tooth, implying only odontoclasts are involved in the semi-basal resorption, which is probably the first step towards evolving a site-specific resorption. The polarized displacement of each generation of resorption surfaces reflects the fact that the cyclic replacement, as well as the sequential addition of tooth sites, is closely related to bone growth. Resorption surfaces and growth arrest surfaces also record the life history and the replacement rate. These data provide unique insights into the origin of osteichthyan tooth replacement.

The role of temperature variation in organismal performance is understudied, but is critically important for understanding the response of biodiversity to climate change. To address this issue in herbivorous insects, I studied the direct and interactive effects of thermal regime (constant vs. fluctuating temperatures) and nutrition (dietary nitrogen) on gypsy moth (Lymantria dispar) performance under laboratory conditions. Predictions for differences between constant and fluctuating thermal conditions were derived from Jensen’s inequality, and artificial diets of differing nutritional quality were made by modifying nitrogen (casein) content. Larvae were reared in the laboratory under four temperature regimes (22°C constant, 22°C fluctuating (±6°C), 28°C constant, and 28°C fluctuating (±6°C)) and two diet treatments (high N, and low N). Gravimetric analyses were also conducted to calculate nutritional indices and assess the short-term effects of temperature and diet quality on fourth instar larvae growth efficiencies. Consistent with predictions from Jensen’s inequality, fluctuating thermal conditions significantly reduced larval performance in both sexes across ontogeny. Low quality diet also reduced performance, but interactions between diet and thermal regime were only found in early instars.

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