Boreal ecosystems are warming roughly twice as fast as the global average, resulting in woody expansion that could further speed up the climate warming. Boreal peatbogs are waterlogged systems that store more than 30% of the global soil carbon. Facilitative effects of shrubs and trees on the establishment of new individuals could increase tree cover with profound consequences for the structure and functioning of boreal peatbogs, carbon sequestration and climate.
We conducted two field experiments in boreal peatbogs to assess the mechanisms that explain tree seedling recruitment and to estimate the strength of positive feedbacks between shrubs and trees. We planted seeds and seedlings of Pinus sylvestris in microsites with contrasting water-tables and woody cover and manipulated both shrub canopy and root competition. We monitored seedling emergence, growth and survival for up to four growing seasons and assessed how seedling responses related to abiotic and biotic conditions.
We found that tree recruitment is more successful in drier topographical microsites with deeper water-tables. On these hummocks, shrubs have both positive and negative effects on tree seedling establishment. Shrub cover improved tree seedling condition, growth and survival during the warmest growing season. In turn, higher tree basal area correlates positively with soil nutrient availability, shrub biomass and abundance of tree juveniles.
Synthesis. Our results suggest that shrubs facilitate tree colonization of peatbogs which further increases shrub growth. These facilitative effects seem to be stronger under warmer conditions suggesting that a higher frequency of warmer and dry summers may lead to stronger positive interactions between shrubs and trees that could eventually facilitate a shift from moss to tree-dominated systems.
Figure 1data_Exp 2
Figure 1 data:
Condition of experimental seedlings in hummocks with contrasting shrub density and tree canopy in Experiment 2: No Trees - Low Shrub biomass (NTLS), No Trees - High Shrub biomass (NTHS), Present Trees - Low Shrub biomass (PTLS) and Present Trees - High shrub biomass (PTHS) during the warmest growing season (2011) and at the end of the experiment (2013). Seedling condition was defined as: healthy (< 50% of the needles turned yellow or brown) or unhealthy (> 50% of the needles turned yellow or brown). Seedlings were 1 month old at plantation time in the July 2010.
Table 1_environmental conditions_Exp 1
Table 1 data:
Environmental conditions and vegetation characteristics in hummocks (circular and bands) and lawns for Experiment 1. Water table depth below surface is an average for the four growing seasons (2010-2013)
Table 2_ photosynthesis data_Exp 1
Table 2 photosynthesis data: Photosynthesis rates for experimental pine seedlings in hummocks (circular and bands) versus adjacent lawns for Experiment 1.
Table 2_seedling responses_Exp 1
Table 2 data: Responses of experimental pine seedlings in hummocks (circular and bands) versus adjacent lawns for Experiment 1 after 4 growing seasons. ST: Seeds inserted on top of moss; SB: Seeds inserted below moss; Small seedling (1 month old at plantation time); Large seedling (2 months old at plantation time). Emergence = % of planted seeds emerged after 1 year. Condition = % healthy seedlings. Stem growth corresponds to vertical stem growth for germinating (ST and SB) seedlings and new stem growth for older (small and large) seedlings.
Table 3_regression seedling-environment_Exp 1
Table 3 data for generalized linear models assessing the responses of experimental pine seedlings in hummocks (circular and bands) and adjacent lawns for Experiment 1 during the whole experimental period (2010-2013). ST: Seedlings from seeds inserted on top of moss; SB: Seedlings from seeds inserted below moss; Small seedling (1 month old at plantation time); Large seedling (2 months old at plantation time). Condition = % healthy seedlings. Growth = stem growth.
Table 4_Environmental data_Exp 2
Table 4: Environmental conditions in hummocks with contrasting shrub density and tree canopy in Experiment 2: No Trees - Low Shrub biomass (NTLS), No Trees - High Shrub biomass (NTHS), Present Trees - Low Shrub biomass (PTLS) and Present Trees - High shrub biomass (PTHS).
Table 4 and Table S5a_seedling performance_Exp 2
Table 4: Seedling performance in hummocks with contrasting shrub density and tree canopy in Experiment 2: No Trees - Low Shrub biomass (NTLS), No Trees - High Shrub biomass (NTHS), Present Trees - Low Shrub biomass (PTLS) and Present Trees - High shrub biomass (PTHS). Seedling emergence, condition and survival from seeds inserted below the moss (SB), and from small planted seedlings.
Table S3_cox regression (survival analysis)_Exp 1
Table S3: Data for Cox survival analysis for experimental pine seedlings in hummocks (circular and bands) versus adjacent lawns during 2010-2013. ST: Seedlings from seeds inserted on top of moss; SB: Seedlings from seeds inserted below moss; Small seedling (1 month old, 10 cm tall at plantation time); Large seedling (2 months old, 30 cm tall at plantation time).
Table S4_ regression seedling-environment 2011_Exp 1
Table S4: Data for generalized linear models assessing the responses of experimental pine seedlings in hummocks (circular and bands) and adjacent lawns for Experiment 1 in 2011. Small seedling (1 month old, 10 cm tall at plantation time); Large seedling (2 months old, 30 cm tall at plantation time). Condition = % healthy seedlings. Growth = stem growth.