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JOURNAL OF GEOPHYSICAL RESEARCH, VOL. 108, NO. D3, 8220, doi:10.1029/2001JD000839, 2003

Annual carbon flux from woody debris for a boreal black spruce fire
chronosequence
B. Bond-Lamberty, C. Wang, and S. T. Gower
Department of Forest Ecology and Management, University of Wisconsin-Madison, Madison, Wisconsin, USA

Received 17 May 2001; revised 7 September 2001; accepted 28 September 2001; published 25 October 2002.
[1] This study examined the distribution and respiration dynamics of woody debris (WD)
in a black spruce-dominated fire chronosequence in northern Manitoba, Canada. The
chronosequence included seven stands that burned between 1870 and 1998; each stand
contained separate well-drained and poorly drained areas. The objectives of this study were
to (1) quantify the distribution of WD, by diameter and decay class, in well-drained and
poorly drained stands across the chronosequence, (2) measure the evolution of CO2 from
WD samples and model the effects of moisture, size and decay on respiration, and (3)
model annual WD respiration and compute decay constants for each site. Coarse woody
debris biomass ranged from 1.4 Mg ha
1 to 177.6 Mg ha
1, generally declining in the older
stands of the chronosequence. More decayed WD had significantly (a = 0.05) higher
moisture, lower density, and higher respiration rates than less decayed WD. Moisture and
decay class were significant predictors of respiration when moisture was below 43%.
Above this level, moisture was not significant, but stand soil drainage was significant, with
drier sites having higher WD respiration. Year of burn was not significant in the respiration
models. Modeled annual carbon emissions from WD ranged from 0.11 to 1.92 Mg C ha
1
yr
1. Modeled annual decay rates, between k = 0.01 and k = 0.06, changed across the age
sequence. Thus a single-exponential decay model may not be appropriate for the age
sequence. Calculating k directly from wood respiration measurements, as done here, may
be useful in allowing examination of year-to-year changes in k. INDEX TERMS: 4806
Oceanography: Biological and Chemical: Carbon cycling; 4815 Oceanography: Biological and Chemical:
Ecosystems, structure and dynamics; 4845 Oceanography: Biological and Chemical: Nutrients and nutrient
cycling; 3322 Meteorology and Atmospheric Dynamics: Land/atmosphere interactions; KEYWORDS: woody
debris, CO2 flux, boreal forest, carbon cycling, black spruce

Citation: Bond-Lamberty, B., C. Wang, and S. T. Gower, Annual carbon flux from woody debris for a boreal black spruce fire
chronosequence, J. Geophys. Res., 107, 8220, doi:10.1029/2001JD000839, 2003. [printed 108(D3), 2003]

1. Introduction studies [Krankina and Harmon, 1995; Stone et al., 1998].

[2] Coarse woody debris (CWD) is a conspicuous and
important characteristic of all forest ecosystems from the
boreal to the tropical. Coarse woody debris influences
nutrient cycling, humus formation, carbon storage, fire
frequency, and water cycling, and serves as a habitat for
both heterotrophic and autotrophic organisms [Harmon et
al., 1986; Rayner and Boddy, 1988]. Dead trees and CWD
have generally not been included in global carbon budgets
[Houghton and Woodwell, 1989; Post et al., 1990], although
their importance as boreal carbon stores have recently
gained attention [Harden et al., 2000; Krankina et al.,
2002; Krankina and Harmon, 1995].
[3] While CWD inventories are relatively straightforward
to perform, measuring decay rates of CWD is difficult due
to physical fragmentation and the slow nature of the decay
process. For these reasons, there are few true long-term
Copyright 2002 by the American Geophysical Union.
0148-0227/02/2001JD000839
One way around this difficulty is to substitute space for time
by employing a chronosequence. A number of studies have
used age sequences to examine CWD distribution in the
boreal forest [Krankina and Harmon, 1995; Lee et al.,
1997; Næsset, 1999; Siitonen et al., 2000; Sturtevant et
al., 1997], and a few have measured respiration rates of
CWD [Chambers et al., 2001; Marra and Edmonds, 1996;
Progar et al., 2000]. We are unaware of any study that has
attempted to use respiration measurements from CWD to
project yearly carbon emissions and decay rates in the
boreal forest, although at least one study has done so for
tropical forests [Chambers et al., 2001].
[4] As CWD is simultaneously a carbon repository, a
source of CO2 emissions, and a wildfire fuel, understanding
its dynamics in boreal forests is particularly important. This
is particularly true for the Canadian boreal forest, where
relatively little research has been done on CWD. More
accurate estimates of CWD respiration and decay rates
should help improve long-term boreal carbon cycling mod-
els that depend on these data [Harden et al., 2000; Harmon

WFX 1-1

WFX 1 - 2 BOND-LAMBERTY ET AL.: CARBON FLUX FOR BLACK SPRUCE WOODY DEBRIS
et al., 1990]. Many climate change scenarios predict a rise
in temperature and fire frequency in the boreal forest [Clark,
1988; Flannigan and Van Wagner, 1991].
[5] The objectives of this study were to (1) quantify the
distribution of woody debris (WD), by size and decay state,
in well-drained and poorly drained black spruce stands
across the chronosequence, (2) measure the evolution of
CO2 from WD samples and model the effects of moisture,
size and decay on respiration, and (3) model annual WD
respiration (RWD) and compute decay constants for each
site. We hypothesized that (1) WD levels would be higher in
well drained areas than in poorly drained ones because of
larger tree sizes in drier areas, and would decrease in the
older stands in the age sequence before rising again with the
increasing contribution of natural tree mortality, (2) more
decayed WD would have higher respiration rates than less
decayed WD, and this respiration would be strongly
dependent on WD moisture content, and (3) when modeled
over the year, WD would be a significant source of CO2
emissions compared to soil respiration.
2. Materials and Methods
2.1. Site Descriptions
[6] The study was conducted in summer 2000 on forest
stands between Thompson and Leaf Rapids, Manitoba,
near the BOREAS Northern Study Area (55 530N, 98
200W). Mean January, July, and annual air temperatures in
Thompson are
25.0, 15.7, and
3.4 C respectively, and
mean annual precipitation is 536 mm; year 2000 was
slightly warmer and drier than these historical averages.
The stands are even-aged and originated from stand-killing
wildfires in 1998, 1995, 1989, 1981, 1964, 1930, and 1870.
The time since burn was 2, 5, 11, 19, 36, 70, and 130
years, res-pectively, at the time of sampling. Separate
poorly drained (wet) and well drained (dry) areas were
identified at each wildfire area; ‘‘wet’’ areas were consid-
ered to be those sites having saturated or poorly aerated
soil and supporting plants adapted specifically to such soil
conditions [Lugo et al., 1990]. Site names used here
accordingly begin with a ‘‘D’’ (dry) or ‘‘W’’ (wet), fol-
lowed by year of burn (e.g., D1998). Most of these stands
were dominated by black spruce (Picea mariana (Mill.)
BSP), although the younger stands also contained jack pine
(Pinus banksiana Lamb.), trembling aspen (Populus trem-
uloides Michx.), white birch (Betula papyrifera Marsh.),
and tamarack (Larix laricina (Du Roi) Koch.). Younger
stands had been black spruce-dominated forests before
burning; the pre-fire forest composition of the two oldest
sites was unknown. The dry sites were on upland, moder-
ately drained clay soils, although the 1995 burn occurred
on kame deposits of sand and gravel. This was the best site
that could be found, given time and logistical constraints.
The wet sites were located on poorly drained clay soils
underlain by discontinuous permafrost.
2.2. Experimental Methods
[7] Downed dead wood (DDW) was defined as woody
debris either lying on the forest floor or standing with a
zenith angle 45 degrees. Because of the small dimensions
of trees (and the woody debris) in the boreal forest, we
collected all DDW 1 cm in diameter. Standing dead trees
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with a zenith angle < 45 degrees were considered standing
dead wood (SDW). The term total woody debris (WD),
defined as DDW + SDW, was used to avoid confusion
with the more standard coarse woody debris (CWD),
which is usually defined as woody debris 10 cm. This
nomenclature is similar to that employed by Idol et al.
[2001].
[8] Surveys were performed at each burn site between
early June and mid-August 2000. Nine 10 m  1 m
transects were established in each of the wildfire areas, four
in well-drained stands and five in poorly drained stands,
respectively. Transects were located at least 200 m from the
edge of a burn to minimize edge effects. The transects in the
well-drained stands were randomly located, and the trans-
ects in the poorly drained stands were oriented end to end
(due to the design considerations of a concurrent study). All
DDW that occurred in the marked transect was collected
and separated by size and decay state in the field; samples
crossing the edge of a transect were cut at the boundary.
Because the study sites had been expressly located in black
spruce stands, most SDW and DDW was black spruce,
although some highly decayed samples were unidentifiable.
Size classes were 1 – 2, 2 –5, 5 – 10, 10– 20, and >20 cm in
diameter. Three decay classes, modeled on Lambert et al.
[1980], were used to classify the WD. These were defined
as follows: class I, a knife could not penetrate into the
sample; class II, a knife could slightly (<0.5 cm) penetrate
the sample, with great resistance; class III, a knife could
fully and with little resistance penetrate the sample. In each
transect, the total DDW in each size/decay class combina-
tion was weighed using a field scale with an accuracy of
50 g. Subsamples were then randomly collected from a few
pieces of DDW and stored in airtight bags for laboratory
analysis. Subsamples collected were as large as possible but
limited by the 33-cm size of the respiration measurement
chamber.
[9] Laboratory analyses were performed within 48
hours. Samples were brought to room temperature (20C)
and WD respiration was measured with a Li-Cor 6200
infrared gas analyzer (Li-Cor Inc., Lincoln, NE). The
analyzer was calibrated daily using 400 ppm CO2 (Scott
Specialty Gases, ±5%, balance N2). Respiration measure-
ments were made in a 33-cm clear acrylic chamber (flow
rate 1100 cm3 s
1, 1 mea-surement every 30 seconds, five
measurements total). This allowed for rapid, accurate
determination of CO2 evolution [Norman et al., 1997].
Wet weight and sample dimensions were recorded, and the
sample was then dried in a forced-air oven at 70C to a
constant weight. Sample moisture (percentage moisture on
a dry mass basis), density (g cm
3), volume, surface area,
and respiration rate (mmol kg
1 s
1) were computed from
the collected data. Volume and surface area were calcu-
lated assuming samples were cylinders, using two diameter
measurements and length.
2.3. Statistical Analyses
[10] Analysis of variance (ANOVA) tests were used to
examine the effects of soil drainage and burn year on WD
biomass and partitioning of biomass among size and
decay classes. The effects of size and decay class on
density, moisture content, and DDW respiration were also
tested.
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BOND-LAMBERTY ET AL.: CARBON FLUX FOR BLACK SPRUCE WOODY DEBRIS WFX 1-3

[11] To develop models for predicting WD respiration, an Table 1. Downed Dead Wood (DDW) and Standing Dead Wood
ANOVA test was used to evaluate the effects of site, soil (SDW) Biomass by Burn Year and Soil Drainage Conditiona
drainage, decay class, size class, and volume to surface area DDW, Mg ha
1 SDW, Mg ha
1
ratio on DDW respiration rates. Because of heteroschedastic Well Poorly Well Poorly
variance, respiration and moisture data were transformed Year of Burn Drained Drained Drained Drained
using natural logarithms. In addition, the respiration-to- 1870 10.5 (8.6) 1.9 (2.4) 0.3 (0.3) 0.5 (0.6)
moisture relationship exhibited an obvious break point, 1930 4.4 (2.5) 1.6 (1.6) 0.6 (0.3) 0.1 (0.1)
related to the fiber saturation point and described below, 1964 13.1 (7.7) 3.6 (1.7) 0.2 (0.3) 0.0 (0.0)
where the linear model changed both slope and intercept. 1981 58.2 (21.6) 14.5 (16.3) 1.3 (1.4) 0.5 (0.2)
An iterative nonlinear model [Chappell, 1989] was used to 1989 177.5 (48.1) 92.0 (23.5) 0.1 (0.3) 2.1 (1.1)
1995 4.1 (2.0) 2.3 (2.3) 3.1 (1.1) 1.0 (0.3)
find the optimum inflection point, minimizing the sum of 1998 19.1 (20.7) 0.4 (0.3) 1.7 (1.1) 1.0 (0.2)
squares error of the two linear models (one above the a
Numbers in parentheses are one standard deviation.
inflection point, and one below). Such a segmented model
allowed us to examine the significance of different effects
above and below the break point, with little loss of fit from a
nonlinear model.
no WD respiration took place. Similarly, in spring, the
[12] Possible independent variable tested in the model-
modeled WD thawed after four weeks of above-zero air
building process included sample moisture, size class, decay
temperatures.
class, wood density, burn year, and soil drainage condition.
[15] The model was iterated on a weekly time step. For
A stepwise elimination procedure was used to remove
each site and decay class, one of two different models could
insignificant terms (at a = 0.05) from the respiration
be used on any specific iteration, depending on WD
models. SAS software version 8.0 [SAS Institute, 1999]
moisture. The default was a ‘‘high-moisture’’ model in
was used for all analyses.
which respiration was only weakly affected by moisture
content. If WD moisture fell below the inflection point at
2.4. Annual Carbon Emission Models
which measurements had shown respiration to be strongly
[13] Annual WD carbon (C) fluxes were computed for the limited by water availability, the low-moisture model was
wet and dry stands at each burn site using models derived used. The respiration rate produced from each model was
from respiration measurements in the lab. Each model scaled using an assumed Q10 of 2.0 (with Q10 = 20.0 below
depended on WD decay class, temperature and moisture 10C). Weekly respiration and C emissions were summed to
throughout the year. Air and soil temperature data were give yearly amounts. The annual decay rate k was calculated
recorded for the entire year using Campbell dataloggers for each burn site as total modeled C respired divided by
installed at each site. Coarse woody debris was assumed to total C present. A C to dry biomass ratio of 0.5 was assumed
follow air temperature on the weekly time step of the model. based on previous studies of Picea and other conifers in
This assumption seems fully justified under the closed Canada [Laiho and Prescott, 1999].
canopies of the older burn sites (1870, 1930, and 1964).
Its validity at the younger burn sites (1981, 1989, 1995, and
1998) is elaborated on in the discussion. Finally, we also did 3. Results
not attempt to model any WD-temperature hysteresis cycle, 3.1. Coarse Woody Debris Distribution
assuming that there was no significant lag for a weekly time [16] Both downed dead wood (DDW) and standing dead
step. wood (SDW) exhibited high variance. Dry mass of DDW
[14] Coarse woody debris moisture data were modeled as ranged from 4.1 to 177.5 Mg ha
1 in the well drained stands,
follows. For each decay class, a separate polynomial model and from 0.4 to 92.0 Mg ha
1 in the poorly drained stands
was fitted to the moisture data observed for the 2000 field (Table 1). Dry mass of SDW in the well and poorly drained
season at all sites. These observed data extended from early black spruce age sequences ranged from 0.1 to 3.1 and 0.0 to
June to mid-August, with time point as the only independ- 2.1 Mg ha
1, respectively. Year of burn (DF = 6/49, F =
ent variable. For all decay classes, these models showed a 76.67, p < 0.0001), drainage class (DF = 1/49, F = 35.62, p <
consistent drying period from week 22 to week 28, followed 0.0001) and their interaction (DF = 6/49, F = 35.62, p <
by an increase in WD moisture until week 33. This pattern 0.0001) had highly significant effects on woody debris (WD)
agreed with the general trend of summer fire danger for mass. Drier stands consistently had more DDW, SDW, and
northern Canada [Stocks and Lynham, 1996], with the WD than wetter ones ( p values for different burn years were
summer 2000 ‘‘Fine Fuel Moisture’’ codes of the Canadian between 0.01 and 0.08). The one exception to this was the
Forest Fire Weather Index (B. Lee, personal communica- 1995 burn site, where WD biomass did not differ signifi-
tion, 2001), and with soil moisture data continuously cantly between the well-drained and poorly drained sites
recorded at all burn sites. It was also much simpler than ( p = 0.27).
precisely modeling the environmental factors controlling [17] In general, decay class I comprised the highest
WD moisture [Brackebusch, 1975; Fosberg, 1971; Harmon percentage of WD (Figure 1). Year of burn had a significant
and Sexton, 1995]. Because of limited WD moisture obser- effect on WD partitioning among decay classes sites
vations in the field, however, we used a single moisture (Table 2). Soil drainage was not a significant term by itself,
model for all sites. Early spring (weeks 18 –21) and fall although the interaction between year of burn and drainage
(weeks 34– 40) were assumed to have high (nonlimiting) class was highly significant. Soil drainage class had no
WD moistures. After the air temperature was <0C for four effect on the partitioning of WD biomass by decay class in
consecutive weeks, WD was assumed to have frozen, and the four oldest (1870, 1930, 1964, and 1981) burn sites. The
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WFX 1 - 4 BOND-LAMBERTY ET AL.: CARBON FLUX FOR BLACK SPRUCE WOODY DEBRIS

Figure 1. Relative distribution (%) of woody debris (WD) mass into size and decay classes by soil
drainage class and year of burn.

1995 and 1964 well drained stands had a significantly
smaller percentage of decay class I material ( p < 0.001)
than other aged stands; the 1995 well drained stand had a
significantly higher percentage of decay class III material
( p < 0.001) than any other stand.
[18] The distribution of WD among size classes was
inconsistent among the different-aged stands for both the
wet and dry areas of the chronosequence (ANOVA table not
shown). Neither year of burn nor soil drainage affected the
size class distribution for size classes 1, 2, or 3. Year of burn,
soil drainage, and their interaction were highly significant
( p < 0.03 for all three effects) for size classes 4 and 5. This was
because the 1989 burn had much larger amounts of size class 4
WD, and the only class 5, than any other year (Figure 1).
3.2. Factors Affecting DDW Respiration, Density,
and Moisture Content
[19] Sample density and moisture content were strongly
affected by decay class. Density of the DDW differed
significantly among the decay classes ( p < 0.001) at each
individual burn site and when data were pooled across the
entire age sequence (Table 3). In addition, DDW moisture
content was significantly lower for decay class I samples
than for decay classes II and III ( p < 0.001). The moisture
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BOND-LAMBERTY ET AL.: CARBON FLUX FOR BLACK SPRUCE WOODY DEBRIS WFX 1-5

Table 2. ANOVA Tables Summarizing the Effects of Burn Year and Soil Drainage on the PARTITION of Woody Debris Biomass Into
Decay Classesa
Decay Class I Decay Class II Decay Class III
Effect DF F p DF F p DF F p
Burn year 6/49 3.18 0.010 6/49 1.86 0.107 6/49 3.19 0.010
Drainage 1/49 1.67 0.203 1/49 3.51 0.067 1/49 0.08 0.785
Burn year  drainage 6/49 4.34 0.001 6/49 4.55 0.001 6/49 2.65 0.026
a
I, least decayed; III, most decayed. For each decay class, the model tested was (percent of biomass in decay class) = burn year + drainage + burn year 
drainage. Degrees of freedom (DF), F, and p values are given.

difference between decay classes II and III was weakly
significant ( p = 0.08).
[20] Sample decay class strongly influenced the measured
respiration rate. The effect of decay class was highly
significant ( p < 0.001), and decay classes I, II, and III
had respiration rates (RWD), averages of 0.16, 0.37, and 0.44
mmol CO2 kg
1 s
1 respectively (Table 3). For samples
from well-drained stands, RWD of decay class I was differ-
ent than decay classes II and III ( p < 0.0001 in each case),
but RWD of decay classes II and III were not different ( p =
0.75). For samples from poorly drained stands, RWD of
decay class III was different than decay classes I and II ( p <
0.01 in each case), but RWD of decay classes I and II was
only weakly different ( p = 0.06).
[21] Sample size class had no effect on DDW density ( p =
0.283), moisture ( p = 0.538), or respiration ( p = 0.143).
When the data were examined by soil drainage class, dry-site
samples still showed no size effect ( p = 0.949), but the wet
site data did show a size effect ( p = 0.003), because the
measured respiration rates for size classes 1 and 2 differed
significantly ( p < 0.001) from size classes 3, 4, and 5.
3.3. Woody Debris Respiration Models
[22] Respiration of WD was strongly correlated with
sample moisture content when moisture was below about
40% (Figure 2). However, at 40– 45% moisture the relation-
ship changed significantly ( p < 0.001 for both slope and
intercept). Optimal model fits resulted from a transition
point at 43.1%. As a result, separate models were developed
for these low-and high-moisture conditions.
[23] Under low (< 43.1%) moisture WD conditions, mois-
ture (DF = 1/95, F = 61.46, p < 0.001) and the interaction of
decay class and moisture (DF = 2/95, F = 4.76, p = 0.011)
were the only significant effects. Burn year, soil drainage,
size, and volume to surface area ratio were not significant.
This model explained only about half of the observed
variation (N = 99, R2 = 0.46, MSE = 0.833). Few decay
class III samples were available at this low moisture level
(Table 4).
[24] The high-moisture model (N = 248, R2 = 0.19, MSE =
0.440) had more significant terms. Moisture (DF = 1/241.
F = 8.33, p = 0.004), decay class (DF = 2/241, F = 3.88, p =
0.022), the interaction of decay class and moisture (DF =
2/241, F = 3.73, p = 0.025), and soil drainage (DF = 1/241,
F = 20.40, p < 0.001) were significant. A contrast test
showed that the well drained 1998 stand (D1998) had a
significantly different intercept, and lower respiration, than
all other burn sites ( p = 0.001). While this difference may
have been due to an initial lag in the decay process [Harmon
et al., 1986], this model is not presented separately due to its
small size (N = 10), and was instead kept in the more general
high-moisture model. Low-moisture models had higher
adjusted R2 values, and more significant moisture terms,
than high-moisture models (Table 4). High-moisture models
for poorly drained sites, and for decay classes II and III WD
from well-drained sites, had R2 ffi 0, i.e., moisture ceased to
be a significant term in the model. This implies that there
was no respiration response to extra moisture for most
samples.
3.4. Annual Carbon Emissions and Decay Rates
[25] The annual model results for a single burn site (1989)
and soil drainage class (well drained) showed the constrain-
ing effects of midsummer drying of WD (Figure 3). Mod-
eled RWD remains roughly constant between weeks 24 and
30, even though average temperature increased by over
10C during this period. Only when WD moisture rises is
the maximum flux (2.82 mmol m
2 s
1 for this stand)
reached. Most other burn sites showed similar moisture
limitations of RWD in midsummer. Modeled carbon emis-
sions at the well drained stands varied between 0.11 and
1.92 Mg C ha
1 yr
1 (Table 5). Carbon emissions from WD
in the poorly drained stands ranged from 0.02 to 1.07 Mg
ha
1 yr
1.
[26] Modeled WD respiration was often significant rela-
tive to the more frequently measured soil respiration. At
D1870 and D1998, RWD comprised 5– 7% of soil respira-
tion (RSOIL) on an annual basis; at the W1870 and W1998
poorly drained sites, the corresponding figure was 1 –2%.
At intermediate-aged burns, however, RWD climbed as high
as 36% of RSOIL in the wet sites, and 54% in dry sites.
[27] Decay rates for WD were calculated to be between
0.016 and 0.045 yr
1 across the well-drained age sequence,
and between 0.022 and 0.032 yr
1 in the poorly drained

Table 3. Average Downed Dead Wood (DDW) Density, Moisture, and CO2 Flux, by Decay and Size Classa
Decay Class Size Class
I II III 1 2 3 4 5
Density, g cm
3 0.44 (0.01) 0.29 (0.02) 0.20 (0.01) 0.36 (0.02) 0.37 (0.02) 0.38 (0.02) 0.39 (0.03) 0.30 (0.04)
Moisture, g g
1 0.44 (0.04) 1.09 (0.09) 1.42 (0.12) 0.75 (0.09) 0.68 (0.07) 0.85 (0.10) 0.63 (0.15) 0.39 (0.07)
CO2 flux, mmol kg
1 s
1 0.16 (0.03) 0.37 (0.04) 0.44 (0.05) 0.22 (0.04) 0.21 (0.03) 0.29 (0.04) 0.35 (0.08) 0.32 (0.12)
a
Numbers in parentheses are one standard deviation.
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WFX 1 - 6 BOND-LAMBERTY ET AL.: CARBON FLUX FOR BLACK SPRUCE WOODY DEBRIS

Figure 2. Woody debris (WD) moisture versus measured CO2 flux, by decay class of CWD sample (I,
least decayed, III, most decayed). The dashed line at 43.1% moisture represents the transition point
between low-and high-moisture models.

stands (Figure 4). For both the dry and wet stands in the age
sequence, decay rates were low at the 1998 burn (0.016 and
0.022 yr
1 respectively), spiked high at the 1995 burn (0.063
and 0.033 yr
1), before rising to a peak at the 36 year-old
1964 burn (0.045 and 0.032 yr
1). Decay rates then declined
in the 1930 and 1870 burns to about (0.025 yr
1).
4. Discussion
4.1. Dynamics of Woody Debris Moisture, Mass,
and Density
[28] The high variability of WD biomass seen here is not
unusual [Harmon et al., 1986] and reflects the spatial
heterogeneity of WD on the landscape. At a number of
burn sites in this study, the standard deviation of WD
biomass is equal to or greater than the mean level of that
site. Many studies of CWD have found a ‘‘U-shaped’’
temporal pattern in debris levels, as initial high levels of
disturbance debris decline and give way first to gradual
input from the regenerating stand, and ultimately to higher
inputs as the mature stand senesces into an uneven age
structure [Lambert et al., 1980; Lee et al., 1997; Spies et al.,
1988; Sturtevant et al., 1997; Pedlar et al., 2002]. The mass
distribution for this age sequence is more variable than this
simple pattern, although there is a minimum at the 1930
burn, i.e., at 70 years. This roughly agrees with Sturtevant et
al. [1997], who found a CWD minimum at about 60 years
for a Abies balsamea-Picea mariana boreal chronosequence

Table 4. Parameter Estimates and Significance for Woody Debris (WD) Respiration Modelsa
Model Parameters for log (RWD) = a + b log (M )
Decay Class N a p b p Adjusted R2 MSE
Low moisture I 77 0.928 0.035 2.641 <0.001 0.50 0.893
Low moisture II 17 0.021 0.969 1.626 0.001 0.52 0.518
Low moisture III 5
0.667 0.659 0.22 0.834
High moisture (well drained) I 25
0.838 0.001 1.300 0.008 0.27 0.724
High moisture (well drained) II 56
0.801 <0.001 0.01 0.455
High moisture (well drained) III 41
0.699 <0.001 0.00 0.270
High moisture (well drained)
High moisture (poorly drained) I 30
1.446 <0.001 0.01 0.552
High moisture (poorly drained) II 59
1.326 <0.001 0.01 0.496
High moisture (poorly drained) III 37
0.879 <0.001 0.00 0.206
a
Models are presented by low (<43.1%) and high (43.1%) WD moisture conditions and by decay class (I, least decayed; III, most decayed). The
dependent variable, WD respiration, is RWD (mmol kg
1 s
1). High-moisture models are further subdivided by soil drainage (well and poorly drained).
Insignificant terms (a = 0.05) are not shown.
 21562202d, 2002, D23, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/2001JD000839 by Maynooth University Library, Wiley Online Library on [20/10/2022]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
BOND-LAMBERTY ET AL.: CARBON FLUX FOR BLACK SPRUCE WOODY DEBRIS WFX 1-7

Figure 3. Temperature, modeled woody debris (WD) moisture, and modeled WD CO2 flux for D1989
(1989 burn, well-drained stand).

in Newfoundland, Canada. But the greater variability seen volatiles other than water in the WD could have distorted the
here, in particular, the high WD biomass levels at the 1989 moisture data, since moisture was computed from the differ-
burn, suggests that the sites in this chronosequence have ence between sample field and dry weights (assuming only
more varied stand and burn histories than expected. water had been volatilized during drying). Previous studies,
[29] It is common for CWD density to decrease as decay however, have found no evidence of gases other than CO2
progresses, and for average moisture to increase [Sollins et released from WD [Boddy, 1983a]. Anaerobic respiration
al., 1987]. Many studies have shown that as CWD moisture occurs at localized sites in decaying wood, and possibly
content falls below about 30%, the fiber saturation point, could have a significant impact in highly saturated WD.
microbes cannot gain access to the remaining water in wood Given the overall modest moisture levels of WD observed in
[Griffin, 1977; Kaarik, 1974; Rayner and Boddy, 1988].
Some researchers have seen a higher level of moisture Table 5. Modeled Maximum Summer Respiration (RWD) From
limitation [Boddy, 1983a; Chambers et al., 2001; Kling- Woody Debris (WD) and Annual Carbon (C) Emissions, by Burn
strom, 1965], and our results also show a relatively high Sitea
transition point, at 43.1% moisture content. This result
assumes a common transition point for all decay classes, Site Maximum Summer C Emissions, Proportion of
RWD, mmol CO2 m
2 s
1 Mg C ha
1 yr
1 Soil C Flux
an assumption that may be unrealistic [Dix and Webster,
D1870 0.18 0.14 5.8
1995]. Respiration can also be depressed at very high levels D1930 0.12 0.11 4.1
of CWD moisture and concomitant oxygen deficiency D1964 0.38 0.30 8.5
[Harmon et al., 1986; Kaarik, 1974]. This has been seen D1981 0.77 0.71 17.3
in the laboratory [Boddy, 1983a; Yoneda, 1980], as well as in D1989 2.82 1.92 53.8
high-rainfall forests, e.g., in the Pacific Northwest [Progar et D1995 0.24 0.22 5.5
D1998 0.22 0.16 7.2
al., 2000]. No such decline was observed in the current W1870 0.04 0.03 1.1
study. Only a few samples were collected with extremely W1930 0.03 0.03 1.1
high moisture levels, however, and the statistical power for W1964 0.07 0.06 2.3
detecting a respiration decline was correspondingly weak. W1981 0.18 0.17 5.7
W1989 1.55 1.07 35.8
4.2. Potential Sources of Error W1995 0.06 0.06 1.5
W1998 0.02 0.02 1.1
[30] Modeling WD respiration flux over the course of a a
Woody debris C flux as a percentage of soil C flux, equal to RWD/RSOIL 
year involved a number of assumptions and potential sources 100, was calculated using data from Wang et al. [2002a]. Sites names
of error. The following errors were probably relatively beginning with ‘‘D’’ are dry (well drained), and those beginning with ‘‘W’’
minor, not greatly affecting the results. The presence of are wet (poorly drained). The ‘‘D’’ or ‘‘W’’ is followed by year of burn.

WFX 1 - 8 BOND-LAMBERTY ET AL.: CARBON FLUX FOR BLACK SPRUCE WOODY DEBRIS
Figure 4. Modeled annual decay rates (k) by years since burn and soil drainage class.
this study, and the much lower rate of anaerobic respiration
in general, we doubt that this effect was significant. In
addition, different species do exhibit different respiration
rates [Kaarik, 1974; Rayner and Boddy, 1988]. But the burn
sites described here are overwhelmingly composed of black
spruce, and almost all identifiable WD was black spruce.
[31] This study was not designed to examine RWD temper-
ature response, and assumed, for all decay classes, a Q10
response to temperature of 2.0 above 10C. This is standard
in many biological systems and is close to values previously
reported for CWD [Boddy, 1983a; Yoneda, 1975]. Raising
the assumed Q10 value by 50%, to 3.0, resulted in an average
decline of 14% in the modeled decay rate; lowering the
assumed Q10 by 25%, to 1.5, resulted in a decay rate increase
of 13% (figures are averages computed from the site-specific
model results). These changes were small because the few
weeks during which temperatures are within 5 of 20C (the
temperature at which sample respiration was measured in the
lab) account for approximately 75% of yearly respiration at
most sites. Thus the model was only moderately sensitive to
Q10 temperature response. A more detailed study of black
spruce WD has found, however, that Q10 does in fact change
substantially as temperature changes [Wang et al., 2002b].
[32] There are a number of other, potentially more serious,
sources of error. Respiration measurements were made in a
33-cm diameter chamber. This put an upper limit on the
sample size that could be measured, restricting our ability to
observe any respiration decline linked to increasing WD
size. The negative correlation between size and decay rate
has sound theoretical underpinnings [Rayner and Boddy,
1988], although it is not always observed in field studies
21562202d, 2002, D23, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/2001JD000839 by Maynooth University Library, Wiley Online Library on [20/10/2022]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
[Harmon et al., 1986; Marra and Edmonds, 1996]. In
addition, many larger-size samples had to be cut to fit into
the measurement chamber. These cut ends could potentially
raise the measured respiration rate by (1) increasing the
available surface area, (2) exposing different-quality sub-
strate to microbial action, (3) increasing gas diffusion in the
interior of the sample, and (4) releasing trapped CO2 in the
interior of the sample. For these reasons, scaling the respi-
ration rates of small cut DDW samples to annual field
estimates could overestimate these annual estimates. Sam-
ples were processed within 48 hours, minimizing the first
three of these effects, but not within six hours, minimizing
the problem of CO2 equilibration [Chambers et al., 2001].
Thus we doubt that these effects were highly significant.
[33] The modeling results also depended on our assump-
tions of WD moisture and temperature regimes. The
assumption that woody debris temperature followed air
temperature seems fully justified under the closed canopies
of the older burn sites (1870, 1930, and 1964), particularly
on the weekly time step employed here. At younger burn
sites, however, sunlight can dramatically heat the upper
surface of WD, even while the lower surface and interior
remain close to air temperature [Graham, 1925; Marra and
Edmonds, 1996]. Because respiration is nonlinear with
temperature, this effect may have led to an underestimate
of WD respiration rates when respiration was not con-
strained by low moisture. Wang et al. [2002b] found that
temperature hysteresis in WD was inversely related to stand
leaf area and that this lag between air and WD temperatures
could have substantial effects on the modeling of WD
respiration.

BOND-LAMBERTY ET AL.: CARBON FLUX FOR BLACK SPRUCE WOODY DEBRIS
[34] Moisture levels in WD were assumed to follow a
model based on field observations, with a midsummer dry
period limiting respiration. As noted above, this assumption
agreed generally with long-term trends, the 2000 Fire
Weather Index, and soil moisture data continuously re-corded
at each stand. For the older closed-canopy sites, in which WD
experiences little direct solar radiation and moderated evap-
orative demand, this model may have overestimated summer
drying and thus underestimated RWD. At a minimum, the
smooth moisture curves used here underestimate the consid-
erable temporal variability seen in WD moisture [Boddy,
1983b] and constitute an unknown source of error. Finally,
the simple freeze-thaw regime employed in the model
undoubtedly masks tremendous variation due to weather,
exposure, etc; but annual respiration was so dominated by
warm-weather production that this effect was probably minor.
4.3. Carbon Emissions and Decay Rates
[35] The annual CO2 flux rates computed here are the
products of a rate term (mmol CO2 kg
1 s
1) and a mass
term (Mg WD ha
1). Because our results showed burn year
having no significant effect on respiration, differences in
site fluxes resulted either from different amounts of WD
biomass (e.g., the biomass peak at the 1989 stand) or
different allocations between decay classes (e.g., the peak
at the 1995 stand, due to high amounts of decay class III
WD). The peak RWD flux occurred 10 –20 years after the
stand-killing fires in both well and poorly drained stands
(Table 5). Additional studies are needed to determine if
these results are an artifact of the exceptionally high WD
biomass levels at the 1989 burn site. However, the pattern is
consistent with reported time required for fire-killed trees to
fall over [Harmon et al., 1986]. Such trees then begin
decomposing at much higher rates [Brackebusch, 1975].
[36] We are aware of only a few studies that have made
concurrent estimates of RSOIL and RWD [Marra and
Edmonds, 1994, 1996], and none that have done so for a
forest chronosequence. Results from this study and Wang et
al. [2002] demonstrate that RWD flux, as a fraction of RSOIL,
varies during succession and can, over the course of the
year, contribute 50% as much CO2 to the atmosphere as
does the soil. If the effects of Q10 and temperature hysteresis
are fully accounted for, the flux may be even higher [Wang
et al., 2002b]. Since soil surface flux is the second-largest
CO2 flux in terrestrial ecosystems [Raich and Schlesinger,
1992], RWD patterns affect a significant component of
landscape CO2 flux.
[37] The very low decay rates k calculated for the 1998
burn (0.016 and 0.022 at the well-and poorly drained stands
respectively) are consistent with an initial time lag in the
decay process [Harmon et al., 1986]. After a spike at the
1995 burn, k has intermediate values at the 1989 and 1981
burns, peaks at the 1964 and 1930 burns, and decreased to
low values in the 1870 burn. The 1995 spike demonstrates
how stand and disturbance history can influence decay
patterns. The large amount of highly decayed pre-burn
(decay class III) WD at this burn site (22.7% overall, versus
1.3% and 2.5% at the 1998 and 1989 burns), ‘‘leftover’’
material from a previous wildfire, seems to have produced
the spike in the decay rate.
[38] The overall average k computed here was 0.031
(0.034 on well drained sites, 0.027 on poorly drained sites).
21562202d, 2002, D23, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/2001JD000839 by Maynooth University Library, Wiley Online Library on [20/10/2022]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
WFX 1-9
This corresponds well with previous studies on Picea abies
in Norway [Næsset, 1999] and Russia [Krankina and Har-
mon, 1995], which found decay rates of 0.033 and 0.034
respectively. It is important to note, however, that the rates
presented here are based only on modeled WD respiration
and ignore fragmentation. As such they are underestimates
in comparison to more traditionally determined k values.
[39] The ‘‘dry’’ stands exhibited higher k values and
higher midsummer respiration rates than the ‘‘wet’’ stands.
Significance tests are not appropriate on model results with
so many unknown sources of error, but these k values were
quite close for most burn sites. The modeled decay rate did
seem to change appreciably with stand age. This may imply
that a multiple-exponential decay model is appropriate for
modeling decay of Picea mariana WD, consistent with the
fact that WD is not homogeneous but rather composed of
different substrates decaying at different rates. The inad-
equacies of a single-exponential model have been noted
before [Laiho and Prescott, 1999; Mattson et al., 1987;
Minderman, 1968; Schowalter, 1992], and such a model
seems inappropriate for this black spruce age sequence,
particularly as not all WD in a stand may originate from the
same disturbance event. If a multiple-exponential model is
indeed needed to adequately estimate WD carbon emissions
at large scales, calculating k directly from wood respiration
measurements, as performed here, may be very useful in
allowing examination of year-to-year changes in k. More
traditional methods do not allow for such a ‘‘snapshot’’ of k
during succession [Harmon et al., 1986].
[40] In conclusion, woody debris was an important car-
bon pool and source of landscape CO2 flux and annual
carbon emissions, and its decay rate was influenced by both
soil drainage and temporal changes. Ecosystem models and
carbon budgets that ignore woody debris may miss a large
component of landscape and regional carbon fluxes, partic-
ularly in regions such as the boreal forest where stand-
killing wildfire is a regular occurrence. An improvement in
our understanding of woody debris distribution and decay
dynamics will allow for better predictions of nutrient and
carbon fluxes from woody debris.
[41] Acknowledgments. This research was supported by grants from
the National Science Foundation (DEB-0077881) and NASA (NAG5-8069)
to S. T. Gower. Ben Bond-Lamberty was supported by an NSF Doctoral
Fellowship. Bryan Lee of Natural Resources Canada generously provided
summer 2000 Canadian Forest Fire Weather Index data.
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B. Bond-Lamberty, S. T. Gower, and C. Wang, Department of Forest
Ecology and Management, University of Wisconsin-Madison, 1630 Linden
Drive, Madison, WI 53706, USA. (bpbond@wisc.edu)