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Linking Restoration and Ecological Succession (Springer ... - Inecol

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54 David A. Wardle <strong>and</strong> Duane A. Peltzer<br />

Table 3.1 Retrogressive successional trends that occur in lake isl<strong>and</strong>s in<br />

northern Sweden as a result of prolonged absence of wildfire. Sources of<br />

information are Wardle et al. (1997, 2003, 2004b) <strong>and</strong> Wardle <strong>and</strong> Zackrisson<br />

(2005).<br />

Response variable<br />

Aboveground:<br />

Trend<br />

Tree vegetation Shift from domination by Pinus sylvestris to<br />

Betula pubescens to Picea abies<br />

Understory vegetation Shift from domination by Vaccinium<br />

myrtillus to Vaccinium vitis-idaea to<br />

Empetrum hermaphroditum<br />

Tree <strong>and</strong> understory biomass Continual decline<br />

NPP Continual decline<br />

Light interception by vegetation Continual decline<br />

Vascular plant diversity Strong increase<br />

Intensity of effects of plant species Continual decline<br />

removals on ecosystem properties<br />

Moss biomass Slight increase<br />

N fixation by mosses Strong increase<br />

Belowground:<br />

Polyphenol concentrations in soil Continual increase<br />

Decomposer microbial biomass Continual decline<br />

Litter decomposition rate Continual decline<br />

Soil carbon sequestration Strong increase<br />

N mineralization rate Continual decline<br />

Mineral N concentration Continual decline<br />

Ratio of mineral N to organic N Continual decline<br />

Ratio of soil N to P Continual increase<br />

The ecological impacts of long-term suppression or absence of wildfire have<br />

been investigated over the past decade through a “natural experiment” involving<br />

forested isl<strong>and</strong>s in Lake Uddjaure <strong>and</strong> Lake Hornavan in northern Sweden<br />

(Wardle et al. 1997, 2003, <strong>and</strong> 2004b, Wardle <strong>and</strong> Zackrisson 2005) (Table 3.1)<br />

(see Fig. 3.3). This system allows significant replication of discrete independent<br />

ecosystems (each isl<strong>and</strong> effectively operates as a separate system) at ecologically<br />

meaningful spatial scales. The main extrinsic driver that varies across<br />

these isl<strong>and</strong>s is wildfire caused by lightning strike; large isl<strong>and</strong>s burn more often<br />

than smaller ones simply because they have a larger area to be intercepted<br />

by lightning. Thus, some of the largest isl<strong>and</strong>s have burned in the past 100 years<br />

while some of the smallest have not burned for 5000 years. These isl<strong>and</strong>s therefore<br />

represent a secondary successional gradient induced by fire history. This<br />

variation across isl<strong>and</strong>s in fire history impacts vegetation composition. Thus,<br />

the largest (most frequently burned) isl<strong>and</strong>s are dominated by relatively rapidgrowing<br />

early successional plant species such as Pinus sylvestris <strong>and</strong> Vaccinium<br />

myrtillus, the middle sized isl<strong>and</strong>s are dominated by Betula pubescens <strong>and</strong> Vaccinium<br />

vitis-idaea, <strong>and</strong> the smallest isl<strong>and</strong>s are dominated by slow-growing,<br />

late successional species such as Picea abies <strong>and</strong> Empetrum hermaphroditum<br />

(Wardle et al. 1997). Picea <strong>and</strong> Empetrum are well known to contain high levels<br />

of secondary metabolites such as polyphenols in their tissues (Nilsson <strong>and</strong><br />

Wardle 2005), so that with increasing time since fire there is a shift from plants<br />

that invest their C in growth to those that invest C in defense. Concomitant with

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