The competitive status of trees determines their
responsiveness to increasing atmospheric humidity – a
climate trend predicted for northern latitudes
ARVO TULLUS
1
, PRIIT KUPPER
1
, ANTS KAASIK
1
, HARDI TULLUS
2
, KRISTA L
~
OH M U S
1
,
ANU S
~
OB E R
1
andARNE SELLIN
1
1
Institute of Ecology and Earth Sciences, University of Tartu, Lai 40, Tartu 51005, Estonia,
2
Institute of Forestry and Rural
Engineering, Estonian University of Life Sciences, Kreutzwaldi 5, Tartu 51014, Estonia
Abstract
The interactive effects of climate variables and tree–tree competition are still insufficiently understood drivers of
forest response to global climate change. Precipitation and air humidity are predicted to rise concurrently at high lati-
tudes of the Northern Hemisphere. We investigated whether the growth response of deciduous trees to elevated air
humidity varies with their competitive status. The study was conducted in seed-originated silver birch and mono-
clonal hybrid aspen stands grown at the free air humidity manipulation (FAHM) experimental site in Estonia, in
which manipulated stands (n = 3 for both species) are exposed to artificially elevated relative air humidity (6–7%
over the ambient level). The study period included three growing seasons during which the stands had reached the
competitive stage (trees were 7 years old in the final year). A significant ‘treatment9competitive status’ interactive
effect on growth was detected in all years in birch (P < 0.01) and in one year in aspen stands (P = 0.015). Competi-
tively advantaged trees were always more strongly affected by elevated humidity. Initially the growth of advantaged
and neutral trees of both species remained significantly suppressed in humidified stands. In the following years,
dominance and elevated humidity had a synergistic positive effect on the growth of birches. Aspens with different
competitive status recovered more uniformly, attaining similar relative growth rates in manipulated and control
stands, but preserved a significantly lower total growth yield due to severe initial growth stress. Disadvantaged trees
of both species were never significantly affected by elevated humidity. Our results suggest that air humidity affects
trees indirectly depending on their social status. Therefore, the response of northern temperate and boreal forests to a
more humid climate in future will likely be modified by competitive relationships among trees, which may poten-
tially affect species composition and cause a need to change forestry practices.
Keywords: Betula pendula, climate change, climate manipulation, elevated air humidity, interactive effects, intraspecific competi-
tion, Populus tremula 9 P. tremuloides, tree–tree competition
Received 6 July 2016; revised version received 7 October 2016 and accepted 14 October 2016
Introduction
Improving our understanding of the impact of climate
change on plant communities requires consideration
of the effects of environmental variables on plant–
plant interaction processes, such as competition (Klan-
derud, 2005; Brooker, 2006; Purves & Pacala, 2008;
Adler et al., 2012; Liancourt et al., 2013). Competition
for resources (primarily for irradiance, water and
nutrients) is a key biotic factor that affects the growth
and functioning of plants at the individual, popula-
tion and community levels (Berger et al., 2008). In for-
ests, competition affects tree mortality, size structure
and species composition (Bugmann & Solomon, 2000;
Reynolds & Ford, 2005; Pretzsch & Sch€ utze, 2014).
However, the interactive effects of climate change
variables and competition in forests are still largely
unknown.
The interactive effects of tree–tree competition and
climate variables have been analysed typically by com-
paring single- and mixed-species plots in climate
manipulation experiments, that is by contrasting the
average population-level responses (Liu et al., 2004;
Kozovits et al., 2005; Kubiske et al., 2007; Smith et al.,
2013). Such studies have demonstrated that trees
responsiveness to climate change factors depends lar-
gely on whether trees are subject to inter- or intraspeci-
fic competition. When growing in mixtures, some
species might have a competitive advantage over
others, thereby changing community species composi-
tion. For instance, the response of Populus tremuloides to
elevated [CO
2
] and [O
3
] differs depending on whether
it is grown in a mixture with Betula papyrifera or Acer
Correspondence: Arvo Tullus, tel. +372 7376 176, fax +372 7376
380, e-mail: arvo.tullus@ut.ee
1961 © 2016 John Wiley & Sons Ltd
Global Change Biology (2017) 23, 1961–1974, doi: 10.1111/gcb.13540