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Litter decomposition rates in Canadian forests

Article  in  Global Change Biology · January 1999

DOI: 10.1046/j.1365-2486.1998.00224.x

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Global Change Biology (1999) 5, 75–82

Litter decomposition rates in Canadian forests

T . R . M O O R E , * J . A . T R O F Y M O W , † B . T AY L O R , ‡ C . P R E S C O T T , § C . C A M I R É , ¶
L. DUSCHENE,** J. FYLES,†† L. KOZAK,‡‡ M. KRANABETTER,§§ I. MORRISON,§
M . S I L T A N E N , ¶ ¶ S . S M I T H , * * * B . T I T U S , † † † S . V I S S E R , ‡ ‡ ‡ R . W E I N § § § and
S . Z O LTA I ¶ ¶
*Department of Geography and Centre for Climate and Global Change Research, McGill University, 805 Sherbrooke St. W.,
Montréal, QC, H3A 2K6, Canada, †Pacific Forestry Centre, Canadian Forest Service, 506 West Burnside Rd., Victoria, BC,
V8Z 1M5, Canada, ‡Taylor Mazier Associates, R.R. #3, St. Andrews, NS, B0H 1XO, Canada, §Faculty of Forestry,
University of British Columbia, Vancouver, BC, Canada, ¶Faculté de Foresterie et Geomatique, Université Laval, Ste. Foy, QC,
Canada, **Great Lakes Forestry Research Centre, Canadian Forestry Service, Sault Ste. Marie, ON, Canada, ††Department of
Natural Resource Science, McGill University, Montreal, QC, Canada, ‡‡Pacific Forestry Centre, Canadian Forest Service,
506 West Burnside Rd., Victoria, BC, V8Z 1M5, Canada, §§B.C. Ministry of Forests, Smithers, BC, Canada,
¶¶Northern Forestry Centre, Canadian Forestry Service, Edmonton, AB, Canada, ***Agriculture Branch, Government of Yukon,
Whitehorse, YK, Canada, †††Newfoundland Forestry Centre, St. John’s, NF, Canada, ‡‡‡Department of Biology, University of
Calgary, Calgary, AB, Canada, §§§Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada

Abstract
The effect of litter quality and climate on the rate of decomposition of plant tissues was
examined by the measurement of mass remaining after 3 years’ exposure of 11 litter
types placed at 18 forest sites across Canada. Amongst sites, mass remaining was strongly
related to mean annual temperature and precipitation and amongst litter types the ratio
of Klason lignin to nitrogen in the initial tissue was the most important litter quality
variable. When combined into a multiple regression, mean annual temperature, mean
annual precipitation and Klason lignin:nitrogen ratio explained 73% of the variance in
mass remaining for all sites and tissues. Using three doubled CO2 GCM climate change
scenarios for four Canadian regions, these relationships were used to predict increases
in decomposition rate of 4–7% of contemporary rates (based on mass remaining after
3 years), because of increased temperature and precipitation. This increase may be
partially offset by evidence that plants growing under elevated atmospheric CO2
concentrations produce litter with high lignin:nitrogen ratios which slows the rate of
decomposition, but this change will be small compared to the increased rate of
decomposition derived from climatic changes.
Keywords: climate change, decomposition, forests, lignin, litter, nitrogen
Received 27 October 1997; revised version received 23 February 1998 and accepted 20 April 1998

Introduction
Decomposing plant litter and soil organic matter play an
important role in the terrestrial global carbon (C) cycle
(Schimel 1995). Of the 1500 Pg (1015 g) of global soil
organic C and 55 Pg C of global litter, about 20 and 70%,
respectively, are found in boreal and temperate forests
(Schlesinger 1997). Global potential net primary produc-
tion in boreal and temperate forests amounts to 13 Pg
annually (Melillo et al. 1993). Litterfall in Canadian forests
is estimated to be 17 Tg (1012) y–1, from a biomass of
12 Pg (Kurz et al. 1992). The decomposing litter eventually
Correspondence: J. A. Trofymow, fax: 11/250-363-0775
e-mail ttrofymow@pfc.forestry.ca
forms soil organic matter and the rate of decomposition
is dependent on environmental controls such as climate,
litter quality and soil characteristics.
Although the influences of climate and litter quality
on litter decomposition rates are well established (e.g.
Meentemeyer 1978, 1984; Melillo et al. 1982; Meentemeyer
& Berg 1986; Taylor et al. 1989, 1991; Aber et al. 1990;
Berg et al. 1993; Aerts 1997), the findings are limited by
the ecological range of sites examined and the small
number of tissue types used. The Canadian Intersite
Decomposition Experiment (CIDET) presents the first
data from a wide range of tissues (tree leaves and needles,
herbs and wood) decomposing at sites ranging from the

© 1999 Blackwell Science Ltd. 75

76 T . R . M O O R E et al.
Fig. 1 The location of the 18 sites and the distribution of ecoclimatic provinces of Canada (after Ecoregions Working Group 1989).
arctic through forests to the prairie border. This study
shows that decomposition rates of 11 litter types over
3 years at 18 sites across Canada can be related to simple
climatic parameters (mean annual temperature and pre-
cipitation) and litter quality, primarily the ratio between
the Klason lignin and nitrogen concentrations (Coûteaux
et al. 1995). These relationships are then used to predict
the effect of climatic change on litter decomposition
rates in Canadian forests, assuming changes in annual
temperature and precipitation from three doubled CO2
General Circulation Model (GCM) scenarios, and changes
in litter quality, resulting from doubled atmospheric CO2
concentrations.
Materials and methods
Rates of litter decomposition were determined at 18
upland forest sites, representing the major ecoclimatic
provinces and regions across Canada (Fig. 1, Table 1).
Details of the study, the sites and methods are contained
in Trofymow (1998). Ten different kinds of litter were
collected in litter traps and from senescent tissues from
a range of forest types (Table 2). In addition, wooden
blocks (10 3 5 3 2 cm) were obtained from a single
western hemlock log, cut to avoid branch knots and
sapwood. The litter was allowed to air-dry, thoroughly
mixed and subsampled to determine the air-dry to oven-
dry conversion factor. Characterization of the litter types
was performed using total elemental analysis (for C, N,
S, P, Ca, Mg and K), wet proximate analysis (for non-
polar extractable, water-soluble, acid-soluble carbo-
hydrates, ash and Klason lignin fractions) and 13C CPMAS
NMR analysis of C fractions on a composite sample that
had been milled to pass through a 0.2 mm mesh. The
methods employed are described in Trofymow et al.
(1995).
Litter bags were made from 20 3 20 cm polypropylene
fabric with openings of 0.25 3 0.5 mm. Bags were filled
© 1999 Blackwell Science Ltd., Global Change Biology, 5, 75–82
 LITTER DECOMPOSITION IN CANADIAN FOREST 77

Table 1 Litter mass remaining, averaged

across all 11 litter types, at 18 upland Mass
sites after 3 years: figure in parentheses
is the standard error of the mean for all Site remaining (%) Latitude Longitude
litter bags collected at each site.
Numerals in bold refer to the site in Chapleau, ON 12 52.8 (4.7) 47°389N 83°149W
Fig. 2. Gander, NF 18 58.2 (6.2) 48°559N 54°349W
Gillam, MN 11 80.1 (4.5) 56°199N 94°519W
Hidden Lake, BC 6 59.1 (4.5) 50°339N 118°509W
Inuvik, NWT 1 86.9 (3.0) 68°199N 133°329W
Kananaskis, AB 7 68.1 (3.9) 51°009N 115°009W
Morgan Arboretum, QC 14 44.1 (4.0) 45°259N 73°579W
Montmorency, QC 15 60.0 (5.7) 47°199N 71°089W
Nelson House, MN 10 79.8 (4.3) 55°559N 98°379W
Petawawa, ON 13 52.6 (3.5) 45°559N 77°359W
Port McNeill, BC 4 46.5 (5.7) 50°369N 127°209W
Prince Albert, SK 9 75.7 (4.1) 53°139N 105°589W
Rocky Harbour, NF 17 52.0 (5.1) 49°329N 57°509W
Schefferville, QC 16 70.4 (5.0) 54°529N 66°399W
Shawnigan, BC 5 43.4 (3.7) 48°389N 123°429W
Termundee, SK 8 71.9 (4.7) 51°509N 104°559W
Topley, BC 3 68.6 (5.0) 54°369N 126°189W
Whitehorse, YK 2 82.3 (3.6) 60°519N 135°129W

Table 2 Litter mass remaining in 11 tissue types, averaged across all 18 sites, after 3 years; figure in parentheses is the standard error
of the mean for all sites and original N content and Klason lignin:N ratio. Letters in bold refer to the tissue types in Fig. 2.

Mass remaining N Klason

Litter type (%) (mg g–1) lignin:N

Aspen leaves (Populus tremuloides) Dpt 56.1 (2.9) 6.7 21.4

Beech leaves (Fagus grandifolia) Dba 70.9 (2.9) 7.1 39.4
Bracken fern (Pteridium aquilinum) Fbf 64.5 (3.2) 8.8 37.4
Black spruce needles (Picea mariana) Csb 56.2 (3.6) 7.3 38.7
Douglas fir needles (Pseudotsuga menziesii) Cdc 67.6 (3.6) 7.0 43.3
Fescue grass (Festuca halii) Gfh 41.8 (1.7) 7.1 15.7
Jack pine needles (Pinus banksiana) Cpj 60.4 (3.5) 12.8 25.6
Tamarack needles (Larix laricina) Cll 66.0 (3.5) 5.9 40.6
White birch leaves (Betula papyrifera) Dbw 49.4 (3.6) 7.2 33.3
Western hemlock wood blocks (Tsuga heterophylla) Whw 88.8 (3.4) 2.4 122.6
Western red cedar needles (Thuja plicata) Ctp 75.3 (3.6) 6.4 55.5

with 10 g of air dry litter, except for those containing
western hemlock wooden blocks, which weighed about
50 g. Litter bags were placed at the field sites in autumn
1992 at four plots at each of the 18 sites. Each litter type
was placed at all sites. The bags were placed on the forest
floor, in contact with the underlying litter; where there
were standing grasses of thick lichen layers, the bags
were placed so that they were in contact with the organic
layers or moss surface. Single bags from the four plots
at each site were collected each year, at the same time as
the original emplacement. After collection, the litter bags
were oven-dried at 55 °C, the litter remaining in the bag
was weighed and the proportion of the original litter
mass calculated. In this paper, we use the data collected
for decomposition after 3 years’ exposure, represented
by the percentage of the original mass that remained.
The data, as quadruplicate samples for each of 11 litter
types at each of 18 sites, are used to identify the effect of
climate (using the mean value of the 11 litter types at each
site) and tissue quality (using the mean value of the 18 sites
for each litter type). Data for each individual bag (n 5 789,
three bags were lost) is then used to identify the combined
effect of climate and litter quality on decomposition rate.
The 3-year data were used because they showed major
differences amongst sites and litter types, and represent
the intermediate stage of decomposition.
Climatic data were collected from the nearest long-
term meteorological station, as the 30-year mean annual
temperature and precipitation. An estimate of annual
actual evapotranspiration for the 18 sites was derived
from the map in the Hydrological Atlas of Canada
(International Hydrologic Decade 1978).

© 1999 Blackwell Science Ltd., Global Change Biology, 5, 75–82

78 T . R . M O O R E et al.

Fig. 2 Relationship between percentage mass remaining after Fig. 3 Relationship between percentage mass remaining after
3 years and mean annual temperature among the 18 forest sites. 3 years and Klason lignin:N ratio among the 11 litter types. The
The line represents the regression from Table 3 and sites are line represents the regression from Table 3 and litter types are
identified by numerals (Table 1). identified by letters (Table 2).

Results and discussion of other chemical properties of the litter (Trofymow et al.
1995), such as Ca, Mg, C and S content, explained , 3.6%
Controls on litter decomposition additional variance each, and NMR characteristics, such
as carboxyl and alkyl C content, explained a further 5
The average litter mass remaining at each site after
and 2%, respectively, to the variance using all samples.
3 years ranged from 43 to 87%, generally increasing from
The observed relationship between decomposition rates
southern to northern sites (Table 1). The relationship
and climate and litter quality variables were combined
between the climatic variables and the mass remaining
into a multiple regression equation, which explained 64%
was established by regressions analysis, using SAS (SAS
of the variance of all litter bags or 73% using the means
Institute 1989). Based on the mean mass remaining at
of replicates (Table 3). The ability of these simple climatic
each site, mean annual temperature and mean annual
and litter quality variables to explain differences in
precipitation accounted for 72–87% of the variance (Fig. 2,
medium-term litter decomposition is very good. How-
Table 3). Mean annual actual evapotranspiration,
ever, about 35% of the variance remains unaccounted for
although significantly related to mass remaining, pro-
and there may be a number of reasons for this, beyond
vided no additional explanation, probably because most
the variation among replicate litter bags which accounts
of these Canadian forest sites experience only limited
for 9% of the variance:
moisture deficit during the summer. A similar relation-
1 During the period of measurement of decomposition
ship was obtained when individual litter bag data (n 5
(1992–95), the annual climate at each site may have
789) were used, although the r2 (0.378) was lower because
differed from the long-term mean used in this analysis.
of the inclusion of variation among litter types and
2 The litter bags were placed on the forest floor or soil
replicates.
surface, where the microclimate will be different from
Tissue quality is of equal importance in determining
that recorded at the nearest meteorological station, and
decomposition rates as climate. The 11 tissue types,
this difference will vary with site.
averaged over all sites, had between 41 and 89% mass
3 Soil conditions, such as acidity, nutrient availability
remaining after 3 years, with the fastest decomposition
and macro- and microbiological activity, will affect the
being observed for the herbs and the slowest for the
rate at which tissues decompose and varies from site
wood block and slowest foliar type for western red cedar
to site.
(Fig. 2, Table 2). This variation can be explained by
4 There may be interaction effects amongst the regression
differences in initial Klason lignin and nitrogen content
variables used. For example, there is a site-specific vari-
in the original tissues (Table 3). The higher the ratio
ation in the slope of the response to Klason lignin:N
between lignin and nitrogen content, the slower the initial
ratio, but this is not easily related to simple climate
rate of decomposition (Fig. 3). The wood block acts as
variables.
an outlier: exclusion of the wood block from the regression
reduces the r2 from 0.74 to 0.65 and increases the regres-
sion slope from 0.39 to 0.71. This may be related to
Effect of global change on litter decomposition rates
the physical characteristics of the block—their size and Climatic change will affect the storage of organic C in
density, compared to the other litter types. Incorporation Canadian forests in a number of ways. Changes in

© 1999 Blackwell Science Ltd., Global Change Biology, 5, 75–82

 LITTER DECOMPOSITION IN CANADIAN FOREST 79

Table 3 Relationship between litter mass

remaining after 3 years and climate and Variable/Regression n r2
litter quality variables: T 5 mean annual
temperature ( °C); P 5 mean annual Climate
precipitation (mm year–1); L 5 lignin Mass 5 65.55(1.88) – 2.39(0.37)T 18 0.722
content (%); N 5 nitrogen content (%); Mass 5 76.32(3.03) – 1.63(0.33)T – 0.015(0.004)P 18 0.865
L/N 5 Klason lignin:N ratio. Figures in Mass 5 76.31(1.35) – 1.63(0.15)T – 0.015(0.002)P 789 0.378
parentheses indicate the standard error
of the regression parameters. Litter quality
Mass 5 53.47(9.85) 1 0.123(0.031)L – 3.34(0.96)N 11 0.750
Mass 5 45.24(3.96) 1 0.392(0.078)L/N 11 0.738
Mass 5 53.40(2.82) 1 0.123(0.009)L – 3.33(0.28)N 789 0.270
Mass 5 45.29(1.18) 1 0.391(0.023)L/N 789 0.265

Combined
Mass 5 59.52(1.24) – 1.63(0.11)T
– 0.015(0.001)P 1 0.390(0.016)L/N 789 0.642
Mass 5 59.44(2.04) – 1.63(0.18)T
– 0.015(0.002)P 1 0.392(0.027)L/N 197 0.729

Table 4 Changes in mean annual temperature ( °C) and precipitation (%) for four forest regions of Canada, based on three doubled
CO2 General Circulation Model scenarios: CCC – Canadian Climate Centre GCM II (Boer et al. 1992); GISS – Goddard Institute for
Space Studies transient GCM (Russell et al. 1995); GFDL – Geophysical Fluid Dynamics Laboratory transient GCM (Manabe et al. 1991).

Model

CCC GIS GFDL

Region °C % °C % °C %

N. Alberta/N. Saskatchewan 1 4.3 1 10 1 1.9 1 13 1 3.3 18

N. Ontario 1 4.6 1 12 1 1.8 16 1 3.5 18
S.E. Ontario/S.W. Québec 1 4.4 14 1 1.5 16 1 3.4 16
Atlantic Provinces 1 4.0 11 1 2.0 15 1 3.4 1 10

temperature and precipitation will affect decomposition
rates, and elevated atmospheric CO2 concentrations may
lead to a change in litter quality. In addition, the net
primary productivity of forests can be expected to change,
as well as the distribution of different forest types.
Predicted changes in mean annual temperature and
annual precipitation for four regions in Canada, derived
from three different doubled CO2 GCM scenarios are
presented in Table 4. In general, these four regions
are likely to experience an increase in mean annual
temperature of between 1.5 and 4.6 °C and an increase
in annual precipitation of between 1 and 12%. These
changes will affect decomposition rates. For example, the
predicted changes in temperature and precipitation from
the CCC and GISS models results in a 3.1–8.5% decrease
after 3 years in the mass of jack pine needles, a common
litterfall component across Canada, compared with the
predicted mass loss using the combined equation in
Table 3. The GISS transient GCM predicts smaller changes
in temperature and precipitation, resulting in smaller
increases in the rates of decomposition compared with
the CCC model, averaging 3.6 and 7.5%, respectively.
Several laboratory experiments have been conducted
on the effect of elevated atmospheric CO2 concentrations
on plants (cf. Ceulemans & Mousseau 1994), including
litter quality, especially nitrogen and, to a lesser extent,
Klason lignin contents (see review by O’Neill & Norby
1996). These results are limited by the short duration of
treatment, the small stature of most of the trees and the
limitations of N availability in the soil (Melillo 1996), and
they show variable responses (Table 5). The studies
suggest, however, that there could be relatively large
decreases in the N content and increases in the Klason
lignin:N ratio of litter, caused primarily by a decrease in
N content. Changes in the Klason lignin:N ratio may affect
decomposition rates, if our finding of the importance of
this ratio in explaining differences in decomposition rate
amongst litter types can be applied to change within a
litter type.
Several studies have found that decomposition rates of
tissues grown at elevated atmospheric CO2 concentrations
are slower than those grown under ambient concentra-
tions: in Lolium roots (Gorissen et al. 1995; Van Ginkel
et al. 1996), in marsh Scirpus (Ball & Drake 1997), and

© 1999 Blackwell Science Ltd., Global Change Biology, 5, 75–82

80 T . R . M O O R E et al.

Table 5 Changes in tissue N and lignin content (%) and lignin:N ratio of plants grown under ambient and elevated atmospheric CO2
concentrations (generally 350 and 500–700 ppm CO2, respectively), expressed as the ratio between elevated and ambient higher CO2
treatments. nd 5 not determined.

Source Vegetation N Lignin Lignin:N

Ball & Drake (1997) Marsh Scirpus and Spartina 1.04–1.10 0.98–1.01 0.98–1.08
Brown (1991) Poplar leaves 0.58–0.75 nd nd
Cotrufo et al. (1994) Deciduous tree leaves 0.67–0.81 1.08–1.21 1.33–1.76
Spruce needles 0.97 1.26 1.30
Cotrufo & Ineson (1995) Birch roots 0.86 nd nd
Ineson (1995) Spruce roots 0.69 nd nd
Coûteaux et al. (1991) Chestnut leaves 0.52 nd nd
Franck et al. (1997) Grass shoots and roots 0.80–1.79 nd nd
Hirschel et al. (1997) Senescent tissues 0.83–1.03 0.90–1.10 0.97–1.22
Johnson (1993) Spruce needles 0.74 nd nd
Kemp et al. (1994) Tallgrass 0.86–1.08 0.94–1.02 0.95–1.09
Luxmoore et al. (1986) Pine needles 1.0 nd nd
Melillo (1996) Deciduous tree leaves 0.62–0.96 1.05–1.08 1.18–1.71
Norby et al. (1986) White oak leaves 0.77 0.64 0.83
O’Neill et al. (1987) Poplar seedlings 0.67 nd nd
Robinson et al. (1997) Grass shoots and roots 0.89–0.93 0.89–1.02 0.96–1.15
Rouhier et al. (1994) Chestnut leaves 0.64 nd nd
Van Ginkel et al. (1996) Grass roots 0.65–0.81 1.12–1.13 1.18–1.72

Table 6 Predicted change (using combined regression in Table 3) in 3-year mass remaining (%) of jack pine needles using predicted
changes in mean annual temperature and precipitation (from Table 4) and an increase in Klason lignin:N ratio (L:N) of 30%, for four
regions in Canada.

Climate change alone Climate change 1 L:N

Region Current CCC GISS CCC GISS

N. Alberta/N. Saskatchewan 68.2 60.7 64.4 63.7 67.4

N. Ontario 61.1 52.6 57.7 55.6 60.7
S.E. Ontario/N.W. Québec 51.0 43.6 47.9 46.6 50.9
Atlantic provinces 47.9 41.3 43.9 44.3 46.9

Betula leaves and roots (Cotrufo et al. 1994; Cotrufo &
Ineson 1995), but there was no significant effect in either
tallgrass prairie (Kemp et al. 1994) or spruce roots (Cotrufo
& Ineson 1995). Robinson et al. (1997) found that decom-
position rates of Festuca shoots and leaves grown at
elevated CO2 were slower than ambient CO2 samples
when placed in a High Arctic site, but the difference was
reversed or insignificant at a Low Arctic site. Franck et al.
(1997) found that the C:N ratio in the litter of four annual
grasses increased when grown under elevated CO2, but
that there was little effect on decomposition rates and
there was strong variability among species. Hirschel et al.
(1997) found little difference in lignin:N ratio of senesced
leaf litter grown under elevated CO2 concentrations,
and there was little difference in decomposition rates,
compared with materials grown under ambient CO2
concentrations. In experiments in which CO2 concentra-
tion has been elevated under field conditions, there is
little evidence of major changes in the N content of litter
(P. Canadell, pers. comm.).
Assuming a 30% increase in Klason lignin:N ratio, the
maximum likely from the laboratory experiments, the
mass remaining of jack pine needles after 3 years’ decom-
position reveals an average reduction of 4.5% for the four
regions with the CC scenario, equivalent to a decomposi-
tion rate increase of 7.9%, compared with present values
(Table 6). With the GISS changes in climate and Klason
lignin:N ratio, the average reduction in mass remaining
is only 0.6, equivalent to an increase of 1.1% in the rate
of decomposition. However, increased rates of soil N
mineralization because of warmer temperatures may
reduce the predicted increase in Klason lignin:N ratios.
Thus, even in the most extreme conditions, changes in
litter quality are unlikely to compensate for increased
decomposition rates associated with warmer and wetter
climatic conditions.

© 1999 Blackwell Science Ltd., Global Change Biology, 5, 75–82

 LITTER DECOMPOSITION IN CANADIAN FOREST
The change in net primary production (NPP) associated
with both warmer and wetter climates and elevated
atmospheric CO2 concentrations is probably of greater
significance to the C budget of forests. Melillo et al.
(1993) used a process-based terrestrial ecosystem model
to predict global changes in NPP for major vegetation
types. They showed that changes in climate and CO2
may lead to increases in NPP of 30–40% in boreal forests
and woodlands, whereas increases were predicted to be
between 5 and 15% in temperate forests. As increased
NPP leads to increased rates of litter fall, this will increase
the store of C both in vegetation and on the forest floor,
especially if the net increases in decomposition rate
are minimal.
Acknowledgements
The study was supported by funding from the Climate Change
Program of the Canadian Forest Service. Technical support was
provided by T. Brown, B. Ferris, R. Leach, E. Andersen, S. Dixon,
J. Kinnis, A. Coneys and A. van Niererk. M. Weber, C. Moreal,
D. Anderson, R. Trowbridge and D. White helped locate sites
or collected initial litter samples. The advice and encouragement
of M. Harmon of the US LTER intersite decomposition study is
also gratefully acknowledged.
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