Land Use, Freshwater Flows and Ecosystem Services in an Era of Global Change

The purpose of this thesis is to analyse interactions between freshwater flows, terrestrial ecosystems and human well-being. Freshwater management and policy has mainly focused on the liquid water part (surface and ground water run off) of the hydrological cycle including aquatic ecosystems. Although of great significance, this thesis shows that such a focus will not be sufficient for coping with freshwater related social-ecological vulnerability. The thesis illustrates that the terrestrial component of the hydrological cycle, reflected in vapour flows (or evapotranspiration), serves multiple functions in the human life-support system. A broader understanding of the interactions between terrestrial systems and freshwater flows is particularly important in light of present widespread land cover change in terrestrial ecosystems. The water vapour flows from continental ecosystems were quantified at a global scale in Paper I of the thesis. It was estimated that in order to sustain the majority of global terrestrial ecosystem services on which humanity depends, an annual water vapour flow of 63 000 km3/yr is needed, including 6800 km3/yr for crop production. In comparison, the annual human withdrawal of liquid water amounts to roughly 4000 km3/yr. A potential conflict between freshwater for future food production and for terrestrial ecosystem services was identified. Human redistribution of water vapour flows as a consequence of long-term land cover change was addressed at both continental (Australia) (Paper II) and global scales (Paper III). It was estimated that the annual vapour flow had decreased by 10% in Australia during the last 200 years. This is due to a decrease in woody vegetation for agricultural production. The reduction in vapour flows has caused severe problems with salinity of soils and rivers. The human-induced alteration of vapour flows was estimated at more than 15 times the volume of human-induced change in liquid water (Paper II).

Interactions between calcium and heavy metals in Norway spruce : Accumulation and binding of metals in wood and bark

Waste products from the forest industry are to be spread in forests in Sweden to counteract nutrient depletion due to whole tree harvesting. This may increase the bioavailability of calcium (Ca) and heavy metals, such as cadmium (Cd), copper (Cu) and zinc (Zn) in forest soils. Heavy metals, like Cd, have already been enriched in forest soils in Sweden, due to deposition of air pollutions, and acidification of forest soils has increased the bioavailability of toxic metals for plant uptake. Changes in the bioavailability of metals may be reflected in altered accumulation of Ca and heavy metals in forest trees, changes in tree growth, including wood formation, and altered tree species composition. This thesis aims at examining: A) if inter- or intra- specific differences in sensitivity to Cd occur in the most common tree species of Sweden, and if so, to study if these can be explained by the uptake and distribution of Cd within the plant: B) how elevated levels of Ca, Cd, Cu and Zn affect the accumulation and attachment of metals in bark and wood, and growth of young Norway spruce (Picea abies): C) how waste products from the forest industry, such as wood ash, influence the contents of Ca, Cd, Cu and Zn in wood and bark of young Norway spruce. Sensitivity to Cd, and its uptake and distribution, in seedlings of Picea abies, Pinus sylvestris and Betula pendula from three regions (southern, central and northern parts) of Sweden, treated with varying concentrations of Cd, were compared. Differences in root sensitivity to Cd both among and within woody species were found and the differences could to some extent be explained by differences in uptake and translocation of Cd. The root sensitivity assays revealed that birch was the least, and spruce the most, sensitive species, both to the external and to tissue levels of Cd. The central ecotype of the species tested tended to be most Cd resistant. The radial distribution, accumulation and attachment of, and interactions between Ca and heavy metals in stems of two-year-old Norway spruce trees treated with elevated levels of Cd, Cu, Zn and/or Ca, were investigated. Further, the influence of these metals on growth, and on root metal content, was examined. Accumulation of the metals was enhanced in wood, bark and/or roots at elevated levels of the metal in question. Even at low levels of the metals, similar to after application of wood ash, an enhanced accumulation was apparent in wood and/or bark, except for Cd. The increased accumulation of Zn and Cu in the stem did not affect the growth. However, Cu decreased the accumulation of Ca in wood. Higher levels of Cu and Cd reduced the stem diameter and the toxic effect was associated with a reduced Ca content in wood. Copper and Cd also decreased the accumulation of Zn in the stem. On the other hand, elevated levels of Ca increased the stem diameter and reduced the accumulation of Cd, Cu, Zn and Mn in wood and/or bark. When metals interacted with each other the firmly bound fraction of the metal reduced was in almost all cases not affected. As an exception, Cd decreased the firmly bound fraction of Zn in the stem. The influence of pellets of wood ash (ash) or a mixture of wood ash and green liquor dregs (ash+GLD), in the amount of 3000 kg ha-1, on the contents of Ca, Cd, Cu and Zn in wood and bark of young Norway spruce in the field was examined. The effect of the treatments on the metal content of bark and wood was larger after 3 years than after 6 years. Treatment with ash+GLD had less effect on the heavy metal content of bark and wood than treatment with ash alone. The ash treatment increased the Cu and Zn content in bark and wood, respectively, after 3 years, and decreased the Ca content of the wood after 6 years. The ash+GLD treatment increased the Ca content of the bark and decreased the Zn content of bark and wood after 3 years. Both treatments reduced, or tended to decrease, the Cd content in wood and bark at both times. To conclude, small changes in the bioavailability of Ca, Cu, Cd and Zn in forest soils, such as after spreading pellets of wood ash or a mixture of wood ash and green liquor dregs from the forest industry, will be reflected in an altered accumulation of metals in wood and bark of Norway spruce. It will not only be reflected in changed accumulation of those metals in which bioavailability in the soil has been enhanced, but also of other metals, probably partly due to interactions between metals. When metals interact the exchangeable bound fraction of the metal reduced is suggested to be the main fraction affected. The small alterations in accumulation of metals should not affect the growth of Norway spruce, especially since the changes in accumulation of metals are low, and further since these decrease over time. However, as an exception, one positive and maybe persistent effect of the waste products is that these may decrease the accumulation of Cd in Norway spruce, which partly may be explained by competition with Ca for uptake, translocation and binding. A decreased accumulation of Cd in Norway spruce will probably affect the trees positively, since Norway spruce is one of the most sensitive species to Cd of the forest trees in Sweden. Thus, spreading of waste products from the forest industry may be a solution to decrease the accumulation of Cd in Norway spruce. In a longer perspective, this will decrease the risk of Cd altering the tree species composition of the forest ecosystem. An elevated bioavailability of Ca in forest soils will, in addition to Cd, probably also decrease the accumulation of other less competitive heavy metals, like Zn and Mn, in the stem.

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.