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421
Understory vegetation as a forest ecosystem
driver: evidence from the northern Swedish
boreal forest
Marie-Charlotte Nilsson1* and David A Wardle1,2
Vegetation research in boreal forests has tended to focus on the tree component, while little attention has been
paid to understory components such as dwarf shrubs, mosses, and reindeer lichens. However, the productivity
of understory vegetation is probably comparable to that of the trees. We review recent research in the boreal
forest of northern Sweden to highlight the ecological importance of understory vegetation, both in the short
term by influencing tree seedling regeneration, and in the longer term by affecting belowground processes
such as decomposition, nutrient flow, and buildup of soil nutrients. Wildfire resulting from lightning strike is
a primary determinant of understory vegetation, and as such is a major driver of forest community and ecosys-
tem properties. Forest management practices that alter the fire regime and the composition of understory veg-
etation may have long-term consequences for both conservation goals and commercial forest productivity.
Front Ecol Environ 2005; 3(8): 421–428
424 Panel 1. Ecological impacts of charcoal from wildfire van Cleve 1986; Zackrisson et al. 1999) and because myc-
orrhizal hyphae produced by ericaceous shrubs directly
Activated carbon contains electrostatically charged surfaces that
adsorb compounds with polar molecules such as phenolics. take up nutrients from recently dead moss tissue, before
Addition of activated C to soil supporting plants that produce the tree seedlings are able to access them (Zackrisson et
these compounds has been shown to reduce their negative al. 1997). This mechanism may also interfere to a lesser
effect on associated plant species (Nilsson and Zackrisson 1992; extent with the growth of previously established, larger
Nilsson 1994; Callaway and Aschehoug 2000).Wildfire produces seedlings and trees, because although their root systems
charcoal, which is a form of activated C, and forest soils in
northern Sweden typically contain between 900 and 2100 kg ha-1 can access nutrients below the depth of mosses and shrub
of charcoal (Zackrisson et al. 1996). Charcoal collected from roots, the moss–shrub layer is able to increasingly immo-
sites that have recently burnt has been shown to remove the bilize available nutrients over time (Wardle et al. 1997;
phenolic batatasin-III (produced by black crowberry) from aque- Zackrisson et al. 1999; DeLuca et al. 2002a). Note that the
ous solutions, and to promote emergence and growth of tree above mechanisms differ from that described by Carleton
seedlings treated with these solutions (Zackrisson et al. 1996).
Charcoal from older fires does not have the same effects and Read (1991) for Canadian boreal forest species, in
because there is increased physical obstruction over time of the which tree seedling mycorrhizae were able to directly
charcoal surface. Furthermore, addition of charcoal to the soil access nutrients from mosses.
surface at levels known to occur in the field has been shown to
strongly stimulate birch seedling productivity and nutrient acqui-
sition for soil collected from under black crowberry (presum- Belowground effects of understory vegetation
ably because of its adsorption of batatasin-III), but not for soil
collected from under other ground-cover species (Wardle et al. The three dominant dwarf shrub species in the Swedish
1998b).The surfaces of fresh charcoal also support high levels of boreal forest differ markedly in their ecophysiological
microbial biomass and activity (Zackrisson et al. 1996; Pieti- attributes. Bilberry has short-lived leaves, grows relatively
käinen et al. 2000) and can support greater rates of ecosystem rapidly, and has poorly defended tissues (ie with low levels
level processes driven by components of the soil microflora,
such as decomposition and nitrification (Zackrisson et al. 1996; of active secondary metabolites), while black crowberry
Wardle et al. 1998b; Pietikäinen et al. 2000; DeLuca et al. 2002b). produces long-lived leaves, usually grows slowly, and has
It has been suggested that the increased dominance of black well defended tissues. The attributes of lingonberry are
crowberry, and associated diminished activity of charcoal with intermediate between these two. Consistent with these
increasing time since burning, contributes to diminished forest species-specific differences, black crowberry produces poor-
tree productivity and biomass, as well as reducing the rates of
those soil processes that promote nutrient supplies available to quality litter compared to that produced by the other
plants (Zackrisson et al. 1996). species. Litterbag studies have shown that black crowberry
litter decomposes more slowly and releases less nitrogen (N)
during decomposition than co-existing ericaceous shrub
leaches into nearby waterways during snowmelt, and con- species (Wardle et al. 2003a,b) and most tree species
centrates in small streams and ponds (Brännäs et al. (Wardle et al. 2003a). Furthermore, a litter-mixing study, in
2004). Short-term experimental studies have shown that which litter from ten over- and understory boreal forest
lethal effects on trout alevins and reduced mobility of species were mixed in all two-way combinations, generally
water fleas (Daphnia spp) were caused by batatasin-III but pointed to black crowberry litter as having the strongest
not by a simpler phenolic compound (Brännäs et al. negative effects out of all species on litter decomposition
2004), thus confirming the specific toxic effect of the rates of associated litter (Wardle et al. 2003b). These below-
bibenzyl structure of batatasin-III (cf. Nilsson et al. 2000). ground effects of black crowberry may impair nutrient sup-
Other understory components in Swedish boreal forests ply rates from the soil and contribute to the adverse effects
also influence tree seedling regeneration and growth, of this species on seedling growth (Nilsson et al. 1999).
although the effects are usually not as strong. For exam- The likely ecosystem-level consequences of dwarf shrub
ple, manipulative field experiments investigating exclu- species are apparent from a study that has been operating
sion of bilberry have shown this species to exert negative for the past 10 years on a series of forested lake islands in
effects on Norway spruce seedlings, although below- northern Sweden (Wardle et al. 1997, 2003a; Wardle and
ground resource competition rather than allelopathy was Zackrisson 2005). These islands represent a retrogressive
probably the mechanism involved (Jäderlund et al. 1997). succession, with different islands representing different
Reindeer lichens do not show negative effects on tree periods of time since last burning. With increasing time
seedlings, and seedlings planted into ground dominated since the most recent fire disturbance, early successional
by lichens show vastly superior growth to those planted species such as bilberry and Scots pine are replaced by
within other ground-layer vegetation (Steijlen et al. later successional species such as black crowberry and
1995; Zackrisson et al. 1995). Meanwhile, seedlings Norway spruce. Domination by black crowberry on these
planted into dense feather moss layers typically establish islands is associated with high concentrations of polyphe-
and grow very poorly, despite the ability of mosses to nolic compounds in the humus, and this in turn appears
retain moisture (Steijlen et al. 1995). There is evidence to contribute to reduced soil microbial activity, lower
that this adverse effect of mosses is because they are very decomposition rates, reduced availability of soil N,
effective in absorbing available nutrients (Oechel and increased soil C sequestration and, ultimately, reduced
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