This study aims to assess biomass and area growth of 600 thalli of the old forest lichen, Lobaria pulmonaria, transplanted to three successional boreal forest stands with (1) natural rainfall regime, (2) additional moistening during dry days, and (3) additional moistening with added nutrients. Mean biomass growth during 100 days varied from 8.3% in the dark young spruce forest to 23.1% in the clear-cut area, with the old forest in between (16.0%). Additional moistening did not enhance lichen growth, probably because the transplantation period was wet. Nutrient additions slightly increased area growth compared to artificial water additions only. Growth was determined by a combination of external (forest stand, site factors) and internal factors (chlorophyll content, biomass per area). Transplants acclimated to high light by increasing thickness and chlorophyll a/b-ratio. Some visible bleaching and a strong positive correlation between chlorophyll content per area and lichen growth in clear-cuts suggest some high light-induced chlorophyll degradation. We believe that biomass growth and natural occurrence of L. pulmonaria is controlled by a delicate balance between light availability and desiccation risk, and that the species is confined to old forests due to a physiological trade-off between growth potential and fatal desiccation damage, both of which increase with increasing light. The discrepancy between potential and realized ecological niches is probably caused by a long-term risk to be killed in open habitats by high light during long periods with no rain.
The amount of large and old trees has decreased in the boreal forests during the last centuries of forestry. Such trees are important habitats for epiphytic lichens and there is a growing concern for lichen species that are associated with large and old trees. However, only little is known about the relative importance of tree size versus age as determinants of lichen diversity. Here we have determined the size, age and growth rate of 157 Norway spruce trees and recorded the occurrence of epiphytic lichen species on their branches and lower stems. The study includes crustose lichens and was done in two old-growth forests in SE Norway. Tree age and tree size were the parameters that explained the largest part of epiphytic lichen diversity. Only the growth rate of the most recent time period, i.e. 1984-2004, showed a statistically significant relationship to diversity. There was no indication of a stabilising species number with increasing tree age. Slow-growing and old trees were, however, mainly of importance to the lichen species growing on stems, and this set of species were in general adversely affected by a large amount of branches. The opposite was the case for the species that were confined to branches as their diversity increased when the amount of branches increased. Our study adds empirical data to support the importance of large and old trees as bearers of biodiversity in boreal forests. Site preservation and patch retention of groups of old and large trees is recommended as measures to maintain epiphytic lichen diversity.
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Nitrogen availability limits growth in most boreal forests. However, parts of the boreal zone receive significant levels of nitrogen deposition. At the same time, forests are fertilized to increase volume growth and carbon sequestration. No matter the source, increasing nitrogen in the boreal forest ecosystem will influence the resource situation for its primary producers, the plants, with possible implications for their defensive chemistry. In general, fertilization reduces phenolic compound concentrations in trees, but existing evidence mainly comes from studies on young plants. Given the role of the phenolic compounds in protection against herbivores and other forest pests, it is important to know if phenolics are reduced with fertilization also in mature trees. The evergreen Norway spruce is long-lived, and it is reasonable that defensive strategies could change from the juvenile to the reproductive and mature phases. In addition, as the needles are kept for several years, defense could also change with needle age. We sampled current and previous year needles from an N fertilization experiment in a Norway spruce forest landscape in south-central Norway to which N had been added annually for 13 years. We analyzed total nitrogen (N) and carbon (C), as well as low-molecular phenolics and condensed tannins. Needles from fertilized trees had higher N than those from controls plots, and fertilization decreased concentrations of many flavonoids, as well as condensed tannins in current year needles. In previous year needles, some stilbenes and condensed tannins were higher in fertilized trees. In control trees, the total phenolic concentration was almost five times as high in previous year needles compared with those from the current year, and there were great compositional differences. Previous year needles contained highest concentrations of acetophenone and stilbenes, while in the current year needles the flavonoids, and especially coumaroyl-astragalins dominated. Condensed tannins did not differ between current and previous year needles from control trees. In conclusion, the phenolic defense of current year needles of mature P.abies trees was strongly changed upon fertilization. This may imply that nitrogen deposition and forest fertilization leave forests less robust in a time when pests may take advantages of a changing climate.
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