Forest Ecology and Management 259 (2010) 2418–2425
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Forest Ecology and Management
journal homepage: www.elsevier.com/locate/foreco
Compositional changes of forest-floor vegetation in young stands of Norway
spruce as an effect of repeated fertilisation
Per-Ola Hedwall
a,∗
, Annika Nordin
b
, Jörg Brunet
a
, Johan Bergh
a
a
Southern Swedish Forest Research Centre, Swedish University of Agricultural Sciences, 230 53 Alnarp, Sweden
b
Umeå Plant Science Centre, Department of Forest Genetics and Plant Physiology, Swedish University of Agricultural Sciences, 901 83 Umeå, Sweden
article info
Article history:
Received 11 January 2010
Received in revised form 18 March 2010
Accepted 22 March 2010
Keywords:
Bryophytes
Ellenberg indicator values
Ground vegetation
Nitrogen
Picea abies
RDA
Vascular plants
abstract
Forestry practices that aim to increase biomass production may mitigate climate change through
increased carbon sequestration and the potential of substituting fossil fuels with renewable biofuels.
Fertilising young stands of Norway spruce in Sweden have shown to increase tree growth by more than
200%. Fertilisation, however, also has other effects on forest ecosystems. Here, we studied the response of
the species composition of forest-floor vegetation to three different frequencies of fertilisation in young
stands of Norway spruce. Fertiliser was applied every year, every second year or every third year. The
total amount of N ranged from 425 kg ha
-1
to 625 kg ha
-1
, in combination with P, K, Ca, Mg, S, Mn, Zi,
B and Cu. The largest effects of the fertilisation were found among bryophytes and lichens, which lost
substantial cover. Unexpectedly, Deschampsia flexuosa, commonly known to be favoured by fertilisation,
was negatively affected. Species that increased in frequency were Oxalis acetosella, Brachythecium sp. and
Plagiothecium sp. Decreased availability of light, as an indirect effect of fertilisation through increased
tree canopy cover, was found to be the most important factor behind the change in species composition
of vascular plants. The total cover of bryophytes, however, did not show any significant response to the
changes in canopy cover, indicating that the effects seen in this group may be a result of more direct
effects of the fertiliser. Few significant differences were found between the two most intensive fertili-
sation frequencies, although fertilisation every third year was often distinguished from both the control
and the other fertilised treatments. Even though the effects at the stand level were substantial, the effects
on biodiversity and function of ecosystems on a landscape or regional level need further investigation.
© 2010 Elsevier B.V. All rights reserved.
1. Introduction
The demands of a growing world population, the need to miti-
gate climate change with increased CO
2
uptake in forests and the
subsequent substitution of fossil fuels with biofuels have recently
led to increased interest in enhancing forest growth through fer-
tilisation (Larsson et al., 2009). In Sweden, fertilisation has been a
common practice in forestry since the 1960s, and a large body of
work exists on the effects of fertilisation on wood production and
the environment (Nohrstedt, 2001; Högbom and Jacobsson, 2002).
Several studies have focused on changes of the forest-floor
vegetation after fertilisation. Commonly recorded effects on vege-
tation due to increased nitrogen levels are an increased abundance
of grasses, such as Deschampsia flexuosa (Kellner and Redbo-
Torstensson, 1995), to the detriment of dwarf-shrubs, such as
∗
Corresponding author. Tel.: +46 40415187; fax: +46 40462325.
E-mail addresses: per-ola.hedwall@ess.slu.se (P.-O. Hedwall),
annika.nordin@genfys.slu.se (A. Nordin), jorg.brunet@ess.slu.se (J. Brunet),
johan.bergh@ess.slu.se (J. Bergh).
Vaccinium myrtillus and V. vitis-idaea (Strengbom et al., 2001, 2002;
Strengbom and Nordin, 2008), although a few studies have also
shown increases of dwarf-shrubs (Kellner, 1993; Mäkipää, 1994).
Bottom-layer species, bryophytes and lichens, normally decrease
in cover after fertilisation (Strengbom et al., 2001; Strengbom and
Nordin, 2008), with species replacement only occurring among
bryophytes (Dirkse and Martakis, 1992).
The effects of fertilisation and clear-cutting of the forest on
forest-floor vegetation resemble each other to a large extent. Clear-
cutting of boreal forest often results in compositional changes in the
forest-floor vegetation. While early succession species like herbs
and grasses increase, species typical of the mature forest, such as
Vaccinium spp. which can be considered boreal keystone species,
decrease in abundance (Bergstedt and Milberg, 2001). The cover of
bryophytes often decreases (Jalonen and Vanha-Majamaa, 2001)
and the proportion of drought resistant species increases (Hannerz
and Hånell, 1997). Even though V. myrtillus is at its lowest abun-
dance following stand-replacing disturbances like clear-cut, this
period has been pointed out as important for the persistence of the
species. Atlegrim and Sjöberg (1996) found that V. myrtillus had as
high vegetative investment in clear-cut areas, as in closed forest,
0378-1127/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2010.03.018