Clines, Species and Eucalypts: An Evolutionary Perspective
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Two eucalypt clines were examined using morphological, ecophysiological and molecular analyses. The species complexes examined were an ironbark complex (Eucalyptus melanophloia x E. whitei) and a box complex (E. brownii x E. populnea). Both of these complexes demonstrate continuous morphological variation across their clines. The origin of these morphological clines has previously been interpreted as the product of secondary contact between allopatric species. In this study, an analysis of morphological variation across the clines did not identify an increase in trait variance in the intermediate populations, which suggests that previous theories concerning the origin of these clines may not be valid. Genetic structuring in nuclear and chloroplast DNA was examined across the clines to investigate whether the morphological clines were the product of secondary contact between two independent evolutionary lineages, or whether the clines represent a single evolutionary lineage that has undergone primary differentiation. The microsatellite analyses indicated that there was little genetic structuring across either cline, and that there were only low levels of population differentiation. The lack of hierarchical structuring in the distribution of nuclear genetic variation suggests that these clines are unlikely to be the product of recent gene flow between two formerly allopatric species/populations. A nested clade analysis of the JLA+ region of the cpDNA provides additional evidence to reject the null hypothesis that the morphospecies classifications represent distinct evolutionary lineages. Instead the analyses indicate that each cline represents a single cohesion species and a single evolutionary lineage. The phylogeographic distribution of cpDNA haplotypes is likely to have resulted from restricted seed mediated gene flow with isolation by distance. A more cogent explanation for the clines, based on the genetic data, is that they have arisen through the process of continuous morphological diversification that has been promoted by a directional selection gradient. Drought experiments were conducted in the glasshouse to investigate whether differences in physiological performance under water stress helps to explain the maintenance of the ironbark cline. Under increasing water stress, the morphotypes showed differences in their ability to maintain water status and photosynthetic rates, yet there was no obvious pattern to these differences across the cline. Physiological differences are therefore inadequate to explain the maintenance of the ironbark cline and highlight the compensatory role that morphological variation may play in alleviating water stress. The value of adopting the cohesion species concept and a hypothesis-testing framework to assess species status is demonstrated in this study. This framework provided a statistical approach to distinguish independent evolutionary lineages from interspecific populations and provides evidence to refute the current species status of the species complexes studied. Eucalypt classification is predominantly based on morphology, which results in taxonomic classification that may not reflect genealogical relationships. This is due to the disparity between morphological and phylogenetic relationships. I therefore suggest that current presumptions regarding the prevalence and importance of hybridisation within the genus may reflect taxonomic classification. An accurate assessment of the prevalence and importance of hybridisation requires species classification to be based on genealogical relationships.
Thesis (PhD Doctorate)
Doctor of Philosophy (PhD)
Australian School of Environmental Studies
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