Changes in tree layer and altitudinal distribution of herbaceous species in temperate old-growth forests over 30 years

Abstract

Temperate forest ecosystems, including old-growth fragments, are subject to increasing pressures, both from biotic and abiotic factors. Frequent disturbance events, rising mean annual temperatures and longer-lasting droughts are causing changes in tree species composition, probably shifting the altitudinal distribution of herbaceous species as well. Our goal was to examine whether such shifts can be observed even in old-growth temperate forests, and if the changes in the species composition and spatial distribution of trees is reflected in the herbaceous layer. Our study was based on a survey of several old-growth forests from the 1970s that was repeated after 30 years. Using spatial point pattern methods and generalized linear mixed effect models, repeated measurements of mapped phytosociological relevés and detailed maps of tree positions from two survey periods allowed us to examine how the species composition of the herb layer and the spatial distribution of trees ≥ 10 cm DBH (diameter at breast height) changed over 30 years. On most of the studied sites, the total number of trees declined and the proportion of broadleaves increased between the two surveys. Analyses of tree spatial distribution showed a general shift from a regular spatial distribution in the 1970s to a clustered spatial distribution of trees in the 2000s. In the 2000s, herbaceous species showed an upwards shift in their distribution compared to the 1970s, even after accounting for the effect of changing tree spatial distributions in both survey periods. These effects could be an outcome of warmer and drier weather conditions during the past decades. Further investigation is needed to examine whether this trend is related to changes in climatic conditions.