A fundamental question of long standing in the study of life on Earth is, “Why are there so many species?” This question concerns the distribution of and relationships among species in the present day, but also requires an understanding of the history of diversity. Patterns of diversity result from multiple, interconnected ecological processes operating at different spatial scales. The goal of this research is to gain knowledge about processes that control diversity by using fossil data to provide a temporal perspective that is unavailable when studying modern ecological communities. The fossil record provides the only natural historical account of changes in the diversity of ecological communities in Earth’s past. This research examines the taxonomic composition and diversity of brachiopod paleocommunities in the Delaware Basin of west Texas (Guadalupe Mountains National Park). The study interval is the Bell Canyon Formation, a 5.4-Myr interval of upper Middle Permian (Capitanian) siliciclastic and carbonate rocks deposited on the toe-ofslope of the basin. Silicified brachiopods extracted from the carbonate rocks provide the basis to test two hypotheses: (1) the taxonomic composition of local fossil brachiopod paleocommunities remains uniform, and (2) the changes in diversity of local fossil brachiopod paleocommunities reflects the relative importance of regional processes. Multivariate analyses of clustering analysis and ordination, diversity partitioning, and rank abundance plots are used to evaluate brachiopod taxonomic composition and diversity within an ecological framework. Sequence stratigraphic analysis provides the means to place the results within an environmental context related to sea-level changes. Results indicate that the reorganization of brachiopod paleocommunity structure coincides with major basinal-scale disruptions. Large disruptions allowed rare taxa and invaders from outside the basin to become dominant within paleocommunities. The dynamics within paleocommunities do not appear to prevent the replacement of the incumbent taxa with new taxa. The importance of these findings indicate that paleocommunities are not static through this interval and can be perturbed into configurations with new dominant taxa. Therefore, ecological responses of paleocommunities are resolvable at the geological time scale.

<p>Elucidating the mechanisms structuring communities has been a challenge for community ecology since its beginnings. One theory argues that assembly rules structure communities by means of deterministic mechanisms arising from biological interactions. Another view maintains that patterns seen in community composition and species abundance result from stochastic processes such as migration and extinction. The dilemma has yet not been resolved unambiguously. The main issue is that communities shaped by deterministic mechanisms can produce stochastic patterns via priority effects. The main goal of this study was to determine whether assembly rules structure communities. My strategy was to minimize priority effects by controlling timing of colonization. To do this I used a null community by combining communities of 17 rock pools. This null community was later divided among experimental communities. I conducted three experiments: (1) Experimental communities were exposed to the same external conditions. (2) Communities were exposed to different environments, disturbance, dispersal and habitat heterogeneity. (3) Replicated null communities were connected to allow inter-replicate dispersal. After 4 months, communities (experiment 1) formed alternative states, suggesting the lack of assembly rules control in community structure. The second experiment showed that adding factors results in more alternative states. The increasing number of alternative states among replicate communities indicates that diversified environment and migration are needed to reproduce qualitative patterns observed in nature. The last experiment (3) showed that patterns observed among connected replicate communities resemble patterns that emerged in the presence of biological interactions in unconnected communities. Similarity of patterns between connected and unconnected groups of communities suggest that local biological interactions can be sufficient to structure communities to a considerable degree. Nevertheless, the regional processes appear necessary in their role of supplying species for local communities.</p> / Doctor of Philosophy (PhD)

The aim of this thesis was to increase knowledge about metapopulation and metacommunity processes in patchy, dynamic landscapes, using epiphytic bryophytes as a model system. Host trees and deciduous forest stands in the coniferous landscape are patchy, temporal and undergo changes in habitat quality during succession. Epiphytes must track this dynamic habitat network for their long-term survival. Community patterns at different spatial scales were explored and linked to regional metapopulation processes and local population dynamics. Spatial structuring in species richness both at a local and regional scale indicated stronger dispersal limitation but lower sensitivity to habitat quality in species with large asexual than in species with small sexual diaspores. In sexually dispersed species, a strong rescue effect was indicated by a bimodal frequency distribution of the species and by increasing local abundance with increasing patch connectivity. Present connectivity to other deciduous forest patches had positive effects on richness of asexually dispersed species, whereas richness of sexually dispersed species was instead related to the landscape connectivity 30 years ago. A study of local growth and reproduction suggested that this is caused by delayed sexual, but not asexual, reproduction. Habitat conditions affected the production of sporophytes, but not of asexual diaspores. No differences in either growth rates or competitive abilities among species with different dispersal and life-history strategies were found. In vitro experiments showed that establishment is higher from large asexual diaspores than from small sexual. Establishment of all diaspore types was limited by pH. There were indications of trade-offs between high germination and protonemal growth rates, desiccation tolerance and a rapid development of shoots from protonema. The results indicated that the epiphyte metacommunity is structured by two main trade-offs: dispersal distance (diaspore size) versus age at first reproduction, and dispersal distance versus sensitivity to habitat quality. Trade-offs in species traits may have evolved as a consequence of conflicting selection pressures imposed by habitat turnover, connectivity and irregular water supply rather than by species interactions. Syndromes of interrelated species traits imply that fairly small changes in habitat conditions can lead to distinct changes in metacommunity diversity: the results indicate that increasing distances among patches cause most harm to asexually dispersed species, whereas cuttings of forests of high age and quality as well as increasing patch dynamics are most harmful to sexually dispersed species. / Senarelagd disputation från: 2009-09-26, Lindahlsalen, EBC, Villavägen 9, 75236 Uppsala, Uppsala, 10:00