Adaptation and Constraint in the Plant Reproductive Phase

Conservatism is a central theme of organismic evolution. Related species share characteristics due to their common ancestry. Some concern have been raised among evolutionary biologists, whether such conservatism is an expression of natural selection or of a constrained ability to adapt. This thesis explores adaptations and constraints within the plant reproductive phase, particularly in relation to the evolution of fleshy fruit types (berries, drupes, etc.) and the seasonal timing of flowering and fruiting. The different studies were arranged along a hierarchy of scale, with general data sets sampled among seed plants at the global scale, through more specific analyses of character evolution within the genus Rhamnus s.l. L. (Rhamnaceae), to descriptive and experimental field studies in a local population of Frangula alnus (Rhamnaceae). Apart from the field study, this thesis is mainly based on comparative methods explicitly incorporating phylogenetic relationships. The comparative study of Rhamnus s.l. species included the reconstruction of phylogenetic hypotheses based on DNA sequences. Among geographically overlapping sister clades, biotic pollination was not correlated with higher species richness when compared to wind pollinated plants. Among woody plants, clades characterized by fleshy fruit types were more species rich than their dry-fruited sister clades, suggesting that the fleshy fruit is a key innovation in woody habitats. Moreover, evolution of fleshy fruits was correlated with a change to more closed (darker) habitats. An independent contrast study within Rhamnus s.l. documented allometric relations between plant and fruit size. As a phylogenetic constraint, allometric effects must be considered weak or non-existent, though, as they did not prevail among different subclades within Rhamnus s.l. Fruit size was correlated with seed size and seed number in F. alnus. This thesis suggests that frugivore selection on fleshy fruit may be important by constraining the upper limits of fruit size, when a plant lineage is colonizing (darker) habitats where larger seed size is adaptive. Phenological correlations with fruit set, dispersal, and seed size in F. alnus, suggested that the evolution of reproductive phenology is constrained by trade-offs and partial interdependences between flowering, fruiting, dispersal, and recruitment phases. Phylogenetic constraints on the evolution of phenology were indicated by a lack of correlation between flowering time and seasonal length within Rhamnus cathartica and F. alnus, respectively. On the other hand, flowering time was correlated with seasonal length among Rhamnus s.l. species. Phenological differences between biotically and wind pollinated angiosperms also suggested adaptive change in reproductive phenology.

Palaeoenvironment in North-Western Romania during the last 15 000 years

The objectives of this thesis are to establish a chronological framework for environmental changes during the last 15,000 years in northwest Romania, to reconstruct the vegetation development, and to evaluate the underlying processes for forest dynamics. Furthermore, an overview of earlier and ongoing pollenstratigraphic work in Romania is provided. Sediments from two former crater lakes, Preluca Tiganului and Steregoiu, situated in the Gutaiului Mountains, on the western extremity of the Eastern Carpathians at 730 m and 790 m a.s.l., respectively were obtained and analysed for high-resolution pollen, macrofossils, charcoal, mineral magnetic parameters and organic matter. The chronostratigraphic framework was provided by dense AMS 14C measurements. Cold and dry climatic conditions are indicated by the occurrence of open vegetation with shrubs and herbs, and cold lake water prior to 14,700 cal. yr BP. The climatic improvement at the beginning of the Lateglacial interstadial (around 14,700 cal. yr BP) is seen by the development of open forests. These were dominated by Pinus and Betula, but contained also new arriving tree taxa, such as Populus, Alnus and Prunus. The gradual establishment of forests may have led to a stabilization of the soils in the catchment. Between ca. 14,100 and 13,800 cal. yr BP the forest density became reduced to stands of Pinus, Betula, Alnus, Larix and Populus trees and grassland expanded, suggesting colder climatic conditions. Picea arrived as a new taxon at around 13,800 cal. yr BP, and between 13,800 and 12,900 cal. yr BP, the surroundings of the sites were predominantly covered by Picea forest. This forest included Betula, Pinus, Alnus, Larix and Populus and, from 13,200 cal. yr BP onwards also Ulmus. At ca. 12,900 cal. yr BP, the forest became significantly reduced and at 12,600 cal. yr BP, a recurrence of open vegetation with stands of Larix, Pinus, Betula, Salix and Alnus is documented, lasting until 11,500 cal. yr BP. This distinct change in vegetation may by taken as a strong decline in temperature and moisture availability. At the transition to the Holocene, at ca. 11,500 cal. yr BP, Pinus, Betula and Larix quickly expanded (from small local stands) and formed open forests, probably as a response to warmer and more humid climatic conditions. At 11,250 cal. yr BP Ulmus and Picea expanded and the landscape became completely forested. The rapid increase of Ulmus and Picea after 11,500 cal. yr BP may suggest the existence of small residual populations close to the study sites during the preceding cold interval. Ulmus was the first and most prominent deciduous taxa in the early Holocene in the Gutaiului Mountains. From ca. 10,750 cal. yr BP onwards Quercus, Tilia, Fraxinus and Acer expanded and Corylus arrived. A highly diverse, predominantly deciduous forest with Ulmus, Quercus, Tilia, Fraxinus, Acer, Corylus and Picea developed between 10,700 and 8200 cal. yr BP, which possibly signifies more continental climatic conditions. The development of a Picea-Corylus dominated forest between 8200 and 5700 cal. yr BP is likely connected to a more humid and cooler climate. The establishment of Carpinus and Fagus was dated to 5750 cal. yr BP and 5200 cal. yr BP, respectively. The dominance of Fagus during the late Holocene, from 4000 cal. yr BP onwards, may have been related to cooler and more humid climatic conditions. First signs of human activities are recorded around 2300 cal. yr BP, but only during the last 300 years did local human impact become significant. The vegetation development recorded in the Gutaiului Mountains during the Lateglacial is very similar to reconstructions based on lowland sites, whereas higher elevation sites seem not to have always experienced visible vegetation changes. The time of tree arrival and expansion during the past 11,500 cal. yr BP seems to have occurred almost synchronously across Romania. The composition of the forests during the Holocene in the Gutaiului Mountains is consistent with that reconstructed at mid-elevation sites, but differs from the forest composition at higher elevations. Important differences between the Gutaiului Mountains and other studied sites in Romania are a low representation of Carpinus and a late and weak human impact. The available data sets for Romania give evidence for the presence of coniferous and cold-tolerant deciduous trees before 14,700 cal. yr BP. Glacial refugia for Ulmus may have occurred in different parts of Romania, whereas the existence of Quercus, Tilia, Corylus and Fraxinus has not been corroborated.