Journal of Ecology 2009, 97, 901912 doi: 10.1111/j.1365-2745.2009.01539.

SPECIAL FEATURE
PLANTSOIL INTERACTIONS AND THE CARBON CYCLE

Soil biota accelerate decomposition in high-elevation

forests by specializing in the breakdown of litter
produced by the plant species above them
Edward Ayres1*, Heidi Steltzer1, Sarah Berg1 and Diana H. Wall1,2
1
Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, CO 80521, USA; and 2Department of
Biology, Colorado State University, Fort Collins, CO 80521, USA

Summary
1. There is mounting evidence that leaf litter typically decomposes more rapidly beneath the plant
species it derived from than beneath the different plant species, which has been called home-eld
advantage (HFA). It has been suggested that this HFA results from the local adaptation of soil
communities to decompose the litter that they encounter most often, which probably comes from
the plant species above them.
2. To test this hypothesis and to investigate how HFA varies over time and in relation to litter
quality, we performed the rst detailed assessment of HFA in relation to litter decomposition. We
monitored decomposition over time in two reciprocal litter transplant experiments involving three
high-elevation tree species that differ in litter quality. The three tree species used were trembling aspen
(Populus tremuloides), lodgepole pine (Pinus contorta) and Engelmann spruce (Picea engelmannii).
3. First, we incubated litter from each of these species with soil biota extracted from stands of
each tree species in a laboratory experiment and observed greater cumulative respiration, a mea-
sure of decomposition, when litter was incubated with its home soil biota. Second, we performed
a eld experiment, which demonstrated that the decomposition HFA also occurred under eld
conditions. In addition, this experiment demonstrated that despite increased mass loss at home,
litter also immobilized more nitrogen when in its home environment. In both experiments, the
HFA was most pronounced for pine litter, which is consistent with the hypothesis that HFA
increases with decreasing litter quality.
4. Synthesis. As well as demonstrating conclusively that soil communities specialize in decomposing
the litter produced by the plant species above them, our data challenge the widely held view that soil
organisms are largely functionally redundant.
Key-words: functional redundancy, home-eld advantage, litter decomposition, reciprocal
transplant, San Juan Mountains, soil biodiversity, subalpine forests

scale a variety of other factors inuence litter decomposition to

Introduction
In terrestrial ecosystems, about 60100% of net primary pro-
duction typically enters the decomposition pathway (Cebrian
1999). As a result of the large amount of material being broken
down, decomposition is an important process in the cycling of
carbon and other nutrients. At the global scale, climate and
substrate quality strongly inuence plant litter decomposition
rates (Aerts 1997; Gholz et al. 2000; Trofymow et al. 2002; Par-
ton et al. 2007; Cornwell et al. 2008). However, at the local
*Correspondence author. E-mail: edayres@nrel.colostate.edu
a lesser degree, including the composition of the soil commu-
nity (Heemsbergen et al. 2004; Wall et al. 2008; Strickland
et al. 2009a), interactions among litter types (Hattenschwiler &
Gasser 2005) and UV radiation (Austin & Vivanco 2006). In
addition, several studies have observed that decomposition fre-
quently occurs more rapidly when litter decomposes beneath
the plant species from which it is derived (i.e. at home) than
beneath different plant species (away) (Bocock et al. 1960;
Hunt et al. 1988; Gholz et al. 2000; Negrete-Yankelevich et al.
2008; Vivanco & Austin 2008), which has been referred to as
home-eld advantage (HFA) (Gholz et al. 2000). However,

 2009 The Authors. Journal compilation  2009 British Ecological Society

902 E. Ayres et al.
HFA is not always observed (Prescott et al. 2000; Chapman &
Koch 2007).
Some authors have speculated that the decomposition-
related HFA results from local adaptation of the soil
community to decompose litter produced by the plant species
above them (Bocock et al. 1960; Wardle 2002; Ayres et al.
2009a). If so, the advantage in HFA would be for the soil biota
(i.e. faster access to the energy and nutrients contained
within the litter). In contrast, plant species may or may not
benet from increased rates of decomposition. If soil com-
munities do locally adapt to the plant species above them,
soil community composition should vary among areas dom-
inated by different plant species, which is a pattern that has
frequently been observed (Grifths et al. 1992; Grayston
et al. 1998; Priha, Hallantie & Smolander 1999; Porazinska
et al. 2003; Bardgett & Walker 2004). Indeed, two recent
incubation experiments reported greater respiration rates, a
measure of decomposition, when soil microbes from herba-
ceous- or forest-dominated ecosystems were incubated with
litter corresponding to their home ecosystem, indicating
that differences in soil community composition among eco-
systems can cause decomposition-related HFA (Strickland
et al. 2009a,b). However, Ayres, Dromph & Bardgett (2006)
tested this hypothesis within a forest ecosystem by incubat-
ing beech, sycamore and sitka spruce leaf litter with soil
biota extracted from stands of each of those tree species,
but found no evidence that litter decomposed faster in the
presence of its home soil community. However, as noted by
the authors, the short duration of the experiment and exclu-
sion of many soil taxa may have limited the ability to detect
HFA (Ayres, Dromph & Bardgett 2006).
A recent re-analysis of litter mass loss data from reciprocal
litter transplant studies reported in the literature indicated that,
on average, 8% more mass was lost from leaf litter when it
decomposed at home than away in forest ecosystems (Ayres
et al. 2009a). There was a large amount of variation in HFA
among reciprocal transplants between different tree species,
ranging from 9% slower decomposition at home than away to
29% faster decomposition at home than away (Ayres et al.
2009a). It is not clear that what factors cause this degree of var-
iability in HFA; however, one factor that could explain some
of the variation is time. This is because, changes in litter mass
remaining during decomposition are often described via a neg-
ative exponential decay model with a constant (k) controlling
the decay rate (e.g. Gholz et al. 2000). Litter decomposition
typically occurs more rapidly at home than away (Ayres et al.
2009a), which suggests that the decomposition constant (k) is
greater for litter decomposing at home than away. Therefore,
if mass remaining of litter decomposing at home (larger k) and
away (smaller k) is plotted on a graph (Fig. 1a), the difference
between litter mass remaining at home versus away represents
HFA (Fig. 1b). Based on this rationale, HFA should increase
with time during the early stages of decomposition, but
decrease with time during the later stages of decomposition
(i.e. when most litter mass has already been lost at home;
Fig. 1b). A similar HFA pattern should emerge with other
measures of decomposition, such as litter N remaining;
 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
(a) (b)
Mass remaining (%)
100
Home-field
advantage
0
Time Time
(c) (d)
N remaining (%)
100
Home-field
advantage
0
Time Time
Fig. 1. Schematic diagram of (a) leaf litter mass remaining at home
(solid line) and away (dotted line). The difference between home and
away litter mass represents (b) the mass loss home-eld advantage.
Similar patterns may occur for litter N loss; however, due to N immo-
bilization during the early stages of decomposition (c), the N loss
home-eld advantage may have an initial negative phase (i.e. where
immobilization occurs earlier at home than away) (d).
although during the earliest stages of decomposition, when N
immobilization often increases with mass loss (Parton et al.
2007), litter decomposing at home may immobilize more
N after a given amount of time than litter decomposing at
an away site (i.e. negative HFA) (Fig. 1c, d). To date, the
relationship between HFA and time has not been investigated
in relation to any measure of decomposition.
Differences in the leaf litter traits could also create differ-
ences in the magnitude of HFA, thereby explaining some of
the variability observed in the literature study by Ayres et al.
(2009a). For example, high-quality litter, which is composed of
compounds that are relatively easy to degrade, might be
expected to have little or no HFA, since most soil communities
will contain biota capable of decomposing these compounds
rapidly. In contrast, low-quality litter, which often contains
highly recalcitrant or toxic compounds, might be expected to
have a much greater HFA, since fewer soil communities will
contain biota capable of degrading these compounds rapidly.
This expectation is supported by the ndings of Hunt et al.
(1988), who performed a reciprocal litter transplant among the
dominant plant species of a forest, mountain meadow and
grassland ecosystem. They found that relatively labile grass
litters from the meadow or the grassland decomposed to a sim-
ilar degree among the three ecosystems, but that recalcitrant
lodgepole pine litter decomposed substantially faster at home
than away (Hunt et al. 1988). Similarly, two recent incubation
studies observed little difference in the decomposition rate of
labile herbaceous litter by different microbial communities,
but found that recalcitrant tree litter decomposed faster in the
presence of microbial communities from forest ecosystems
than herbaceous ecosystems (Strickland et al. 2009a,b).
To investigate HFA in relation to leaf litter decomposition,
we performed reciprocal litter transplants among the three
dominant tree species (trembling aspen, lodgepole pine and
Engelmann spruce) in a high-elevation ecosystem in southwest
 Home-eld advantage hastens decomposition 903

Colorado, USA. First, we established a laboratory incubation elevation site was used in the laboratory experiment (see below), we
experiment to test the hypotheses that (i) differences in soil assessed litter quality separately at this site using values from each
biotic community composition associated with different plant subsample and set litter type as the only main effect. In all cases, data
species result in litter decomposition HFA; (ii) HFA exhibits a were log-transformed to meet assumptions of normality and homoge-
neity of variance.
humpback (i.e. sinusoidal) relationship over time; and (iii) the
magnitude of HFA will increase with decreasing litter quality.
Secondly, we established a eld-based reciprocal litter trans- LABORATORY EXPERIMENT
plant experiment to determine whether HFA also occurred The laboratory experiment focused on the lowest elevation site of the
under eld conditions and to investigate impacts on nutrient four eld sites we identied (3742.595N, 10745.112W). Eighty soil
cycling. Specically, we hypothesized that as well as increasing cores (3.4 cm diameter, 10 cm deep) were collected from the stand of
litter mass loss and respiration rates, litter nitrogen (N) release each tree species at this site in October 2007 (i.e. 240 cores in total).
and soil inorganic N concentrations below the litter would be Twenty of these soil cores from each stand were sieved (2 mm) to
lower than expected when litter was at home during the early remove roots and rocks. This sieved soil from each stand was subse-
stages of decomposition, but higher than expected during later quently mixed together to create a single soil substrate and was steril-
stages of decomposition (based on Fig. 1c, d). ized (autoclaved at 121 C for 30 min). This sterile soil was separated
into three subsamples, which were sequentially inoculated with
microbes, microfauna and mesofauna extracted from the remaining
Materials and methods soil cores. One of the subsamples of the sterile soil was inoculated
with soil biota extracted from aspen soil, another with biota extracted
SITE DESCRIPTION from pine soil, and the last with biota extracted from spruce soil. This
resulted in three soils that were identical in every respect, except for
Our study focused on high-elevation (3000 m a.s.l.) forest ecosys-
differences in their soil biotic community, which corresponded to the
tems 914 km south-west of Silverton in the San Juan Mountains,
soil community associated with aspen, pine or spruce stands.
Colorado, USA. Trembling aspen (Populus tremuloides), lodgepole
To inoculate the sterile soil with microbes, 20 soil cores from each
pine (Pinus contorta) and Engelmann spruce (Picea engelmannii) are
stand were separately crumbled and each core was mixed with
dominant tree species in this region. These trees often occur in almost
150 mL water, stirred vigorously and left to stand for 1 h, stirred
pure stands, with stands of the different species frequently adjacent
again before the liquid fraction was poured through a 100-lm screen.
to one another. We identied four sites where near-pure stands of
The liquid fraction was then poured onto the sterile soil subsamples.
aspen, pine and spruce occurred in close proximity to one
Microfauna were extracted from 20 soil cores from each stand using
another (3742.595N, 10745.112W; 3742.373N, 10744.785W;
Baermann funnels (Coleman et al. 1999) and added to the soil.
3742.974N, 10744.633W; and 3743.895N, 10741.905W). The
Mesofauna were extracted from the remaining 20 soil cores from each
shortest and longest distance between any two sites was 1 and 5 km,
stand using Tullgren funnels (Coleman et al. 1999) and were collected
respectively. To minimize microsite differences among the stands
on moistened cotton balls, which were subsequently placed on the soil
within a site (other than those directly caused by the tree species), we
surface. The inoculation with soil biota occurred over a 4-week period:
ensured that the stand of each tree species within a site occurred at a
microbes were added to the soil over the rst 2 weeks, mesofauna were
similar elevation, aspect, slope and on the same bedrock. The short
added during weeks 2 and 3, and mesofauna were added during weeks
distance between stands (<200 m) within a site ensured that each
3 and 4. During this time the soils were sealed in containers to prevent
stand had a similar macroclimate. The sampling sites were located
colonization by airborne microbes. Similar re-inoculation procedures
within an area that was extensively burned in 1879 (Jamieson, Rom-
have been performed in other studies (Laakso & Setala 1999; Grifths
me & Somers 1996). Forests returned after the re, but the US Forest
et al. 2001; Ayres et al. 2004; Heath et al. 2005), but it is probable that
Service also planted lodgepole pine trees, which were not previously
rare species and species intolerant of disturbance are underrepresented
present in this region, to quicken the development of a forested eco-
in re-inoculated communities. Following the inoculation process, the
system (Paulson & Baker 2006).
re-inoculated soils were incubated at room temperature for 3 weeks,
In late August 2006, litter traps consisting of window screen mesh
watered as necessary to prevent drying, and mixed weekly to ensure
were placed in stands of the three tree species at each of the four sites
colonization throughout the soil. During this time, fungal hyphae
to collect naturally senesced leaf litter. Leaf litter was collected from
growth and soil mesofauna activity could be seen on the soil surface,
litter traps in early October 2006, sorted to remove non-target litter
indicating that the inoculation process was successful.
and air-dried. Three subsamples of air-dried litter from each species
The re-inoculated soil was gently mixed and 20 g (dry weight equiv-
at each site were oven-dried (65 C) to determine initial moisture con-
alent) was placed into sixty 120-mL specimen cups (20 cups each
tent, while total C and N content was determined using an elemental
received soil inoculated with aspen, pine or spruce biota). Soil mois-
analyzer (LECO, St Joseph, MI, USA) and lignin concentrations
ture was increased to 60% water-lled pore space (52% moisture con-
were determined using the forest products method (Ryan, Melillo &
tent). One gram of air-dried leaf litter was placed on the soil surface of
Ricca 1990). In addition, an assessment of soil biotic community
45 soil-lled specimen cups; the remaining 15 specimen cups were used
composition in the stands of aspen, pine and spruce in October 2006
for soil-only treatments. Fifteen specimen cups received aspen litter,
revealed differences in the abundances of mesostigmatid mites, coll-
15 received pine litter and the remaining 15 received spruce litter. As a
embolans and rotifer, and in fungal, bacterial and nematode commu-
result, leaf litter from each tree species was incubated with soil biota
nity composition (Ayres et al. 2009b).
extracted from beneath each tree species in all possible combinations,
Litter quality (%N and lignin:N) data were analysed using anovas.
with each treatment replicated ve times. Five additional specimen
To assess litter quality across all four sites the mean of the three subs-
cups received neither soil nor litter and acted as controls. The speci-
amples of each litter type at each site was used and the anova included
men cups were placed in air-tight mason jars so that respiration,
litter type and block (site) as main effects. As only litter from the low

 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
904 E. Ayres et al.
a measure of decomposition, could be determined over time. A repli-
cate of each treatment was placed in one of ve cardboard boxes,
which corresponded to the block effect in the statistical analysis. A
small amount of water was placed in each mason jar to maintain a
humid atmosphere and the jars were incubated at 20 C, which was
approximately equivalent to soil temperatures on a hot summer day.
Carbon dioxide concentrations in samples of headspace air, sampled
via rubber septa, were determined using an infrared gas analyzer
(LI-COR, Lincoln, NE, USA). Mason jars were vented whenever
CO2 concentrations exceeded 20 000 ppm in any jar. Initially, head-
space samples were taken daily, but sampling frequency was gradually
reduced over the course of the experiment. The experiment is ongoing,
but results are presented from the rst 225 days. Based on mass loss of
the same litter species in a similar incubation experiment (unpublished
data), litter mass loss after 225 days in the laboratory experiment
should be similar to mass loss observed at the end of the 2-year eld
experiment.
Cumulative respiration data were analysed using a repeated-mea-
sures anova including main effects and interactions among litter type
(aspen, pine, spruce or no litter), soil biotic community (aspen, pine or
spruce), day (repeated-measure), and block (replicate). Where signi-
cant effects of litter type or soil biotic community were observed, Tukey
HSD tests were used to determine which treatments differed signi-
cantly (P < 0.05) from one another. See the section on Calculating
home-eld advantage for details on how HFA data were analysed.
FIELD EXPERIMENT
In October 2006, 3 g of air-dried aspen, pine or spruce litter from each
site was placed into 54 litterbags (10 10 cm, 1 mm mesh), i.e. 18
litterbags per litter species per site. Since there were four sites, this
resulted in a total of 216 litterbags (3 litter species 4 sites 18 litter
bags = 216 litterbags). The litterbags were placed in the stands of
aspen, pine and spruce in a reciprocal litter transplant design, so that
each stand received 18 litterbags (six of each litter species to allow for
multiple sampling dates) and litter collected from each of the four
sites was only returned to the stands within that site (for instance,
litter collected from the lowest elevation site was only returned to the
stands at the low elevation site). The litterbags were deployed on 78
October 2006 and freshly fallen litter was removed from the surface
of the forest oor prior to place the litterbags on the ground (the rest
of the litter layer was not disturbed). The six litterbags of each litter
type within a stand were placed in a ring around a 10-cm diameter,
5-cm high plastic collar, which was used for in situ respiration mea-
surements. The plastic collar was inserted into the soil and 3 g of air-
dried litter was placed in and around the collar (litter type corre-
sponded to the litter in the surrounding litterbags). The litterbags and
plastic collar were covered by a coarse plastic mesh (1 1 cm holes)
to minimize disturbance of litterbags by animals. In addition, one
iButton temperature data logger (Maxim Integrated Products, Sun-
nyvale, CA, USA) was placed in the surface soil (3 cm deep) of each
stand and set to record temperature at 4-h intervals.
One litterbag of each litter type in each stand was collected over ve
sampling dates: 1415 May 2007 (day 220); 2324 July 2007 (day
290); 2425 September 2007 (day 353); 1011 June 2008 (day 612);
and 910 October 2008 (day 734); the remaining litterbag acted as a
spare. Thus, the experiment design was: 3 litter species 3 stands spe-
cies 4 sites 5 sampling dates + 36 spare litterbags = 216 litter-
bags. Each year, the early-season sampling date corresponded to the
sites becoming snow free and the late-season sampling date corre-
sponded to anticipated snowpack development. In situ soil respira-
tion rates were measured at each sampling date using an infrared gas
 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
analyzer (PP Systems, Amesbury, MA, USA) placed on the plastic
collar (i.e. measuring combined heterotrophic and root respiration
from the litter and soil layer). Respiration was not measured at the
last sampling date due to inclement weather. Following the collection
of a litterbag, a soil core (3.4 cm diameter, 10 cm deep) was taken
directly beneath the centre of the litterbag to assess soil inorganic N
concentrations. Soil nitrate and ammonium concentrations were
determined via potassium chloride extraction and analysed on an
autoanalyser (OI Analytical, College Station, TX, USA) and
expressed on a soil dry weight basis (soil was oven-dried at 105 C to
determine moisture content). Litter was oven-dried (65 C) and
weighed to determine mass loss and subsequently ground for total C
and N analysis on an elemental analyser (LECO, St Joseph, MI,
USA).
Litter mass remaining, litter N remaining, soil and litter respiration
rates, soil inorganic N, and soil moisture data were analysed using
anovas (a repeated-measures anova was used for respiration data).
The anovas included litter type (aspen, pine or spruce), stand (aspen,
pine or spruce) species, day (repeated-measure for respiration data)
and block (site) as main effects, as well as their interactions. Soil inor-
ganic N data were log-transformed to meet assumptions of normality
and homogeneity of variance. Where signicant effects of litter type
or stand species were observed, Tukey HSD tests were used to deter-
mine which treatments differed signicantly (P < 0.05) from one
another. See the section on Calculating home-eld advantage for
details on how HFA data were analysed. Mean monthly soil tempera-
tures were calculated for each time of day (3am, 7am, 11am, 3pm, 7pm,
11pm; Mountain Standard Time) and analysed using a repeated-mea-
sures anova, which included stand species, block (site), time of day
and month (repeated-measure) as main effects, as well as their inter-
actions.
CALCULATING HOME-FIELD ADVANTAGE
Calculating HFA is more complex than simply comparing the decom-
position rates for a litter type at its home site and an away site,
because differences in environmental conditions (e.g. microclimate or
nutrient availability) between the sites could also inuence decompo-
sition. Previous reciprocal litter transplant studies that have
attempted to identify HFA have focused on the interaction between
site and litter type. If this interaction does not signicantly inuence
the measure of decomposition, then there is no HFA; however, if
there is a signicant interaction, it may indicate HFA, but not neces-
sarily. In addition to this ambiguity, simply identifying a statistically
signicant HFA does not provide information on its magnitude,
which is probably more relevant.
Here, we calculated litter decomposition HFA using a method
originally developed to calculate HFA in sports (Clarke & Norman
1995). This method allows an HFA value to be calculated for each
species involved in a reciprocal transplant and it is expressed in the
units of the measurement (e.g. % mass loss or g CO2 m)2 h)1).
Home-eld advantage is calculated using the following equations
(adapted from Clarke & Norman 1995):
ADHi HDDi  ADDi  H eqn 1
HDDi DiI  DjI DiI  DkI eqn 2
ADDi DiJ  DjJ DiK  DkK eqn 3
H HDDi HDDj HDDk =N  1 eqn 4
where ADHi is the additional decomposition at home for species
 Home-eld advantage hastens decomposition 905

i; i, j and k are litters derived from plant species i, j and k, Table 1. P values from anovas of cumulative respiration and
respectively; I, J and K are areas dominated by species i, j and k, additional cumulative respiration at home (ADH in eqn 1) from the
respectively; D is a measure of decomposition (e.g. litter mass loss laboratory experiment. The biota treatment does not apply to the
or respiration); HDD and ADD represent home decomposition additional respiration values. Note that P values relating to
difference and away decomposition difference, respectively; additional cumulative respiration at home refer to differences among
H represents the total HFA for all species combined; and N rep- treatments, not whether there is a signicant HFA or not. Asterisks
resents the number of species. If ADHi > 0, litter from species i on the corresponding graphs (Fig. 2) indicate whether there is a
decomposed faster than expected when at home (i.e. HFA); if signicant HFA (i.e. whether the data is signicantly different from
ADHi = 0, litter decomposition at home occurred at the rate zero)
that was expected (i.e. no HFA); and if ADHi < 0, litter decom-
position at home occurred slower than expected (i.e. home-eld Cumulative Additional cumulative
disadvantage). We calculated ADH for each species in each block respiration respiration at home
(replicate in the laboratory experiment and site in the eld experi-
Litter <0.001 <0.001
ment). anovas were performed on the resulting values to test for
Biota 0.002
signicant (P < 0.05) differences among treatments. The anovas
Day <0.001 <0.001
included species (aspen, pine or spruce), day (which was a Litter biota 0.490
repeated-measure for the analysis of respiration data), and block Litter day <0.001 <0.001
(replicate in the laboratory experiment and site in the eld experi- Biota day <0.001
ment) as main effects, as well as their interactions. Soil inorganic Litter biota day <0.001
N data were log-transformed to meet assumptions of normality Block 0.030 0.001
and homogeneity of variance. We used t-tests to determine Litter block 0.948 <0.001
whether the ADH values from equation 1 differed signicantly Biota block 0.593
(P < 0.05) from zero (i.e. whether HFA occurred). To limit the Day block 0.005 0.007
Litter biota block <0.001
number of t-tests that were performed, data were combined
Litter day block 0.001
across sets of ve sampling dates.
Biota day block 0.508

Results
LITTER QUALITY
Litter N across all sites was 0.84 0.08, 0.47 0.03 and
0.41 0.01% (mean SE) for aspen, pine and spruce,
respectively (species P < 0.001; block P = 0.808), whereas
lignin:N was 24 1, 73 6 and 47 2 (mean SE) for
aspen, pine and spruce, respectively (species P < 0.001; block
P = 0.570). At the lowest elevation site only (i.e. the litter used
in the laboratory experiment), litter N was 0.74 0.03,
0.56 0.02 and 0.41 0.01% (mean SE) for aspen, pine
and spruce, respectively (species P < 0.001), whereas lignin:N
was 27 0, 61 2 and 51 2 (mean SE) for aspen,
pine and spruce, respectively (species P < 0.001).
LABORATORY EXPERIMENT
Since the laboratory experiment is currently ongoing, we are
unable to directly assess microbial and faunal community com-
position in each re-inoculated soil as this would severely dis-
turb the samples. However, cumulative respiration from soil
re-inoculated with biota from aspen, pine, or spruce stands dif-
fered signicantly from one another (Table 1, Fig. 2; see
Fig. S1 for all combinations of litter and soil biota), indicating
that we successfully established distinct soil communities dur-
ing the inoculation process.
Substantially, greater amounts of CO2 were respired from
microcosms that contained leaf litter and soil than from soil-
only microcosms indicating that most CO2 was derived from
respired litter (Fig. 2a; Fig. S1), although some of this addi-
tional CO2 may have resulted from a priming effect on soil car-
bon (Dijkstra & Cheng 2007). Cumulative respiration was
greater from microcosms containing aspen litter than pine or
spruce litter (Table 1, Fig. 2a) and from microcosms contain-
ing aspen or pine soil biota than spruce soil biota (Table 1,
Fig. 2b). There was a signicant interaction between litter, soil
biota and day on cumulative respiration, indicating a potential
for HFA (Table 1). Indeed, there was signicantly greater res-
piration when litter was incubated with its home soil biota for
all litter types combined, but this effect varied with time being
signicantly greater than zero from day 0 to 7 and day 97 to
225 (Fig. 2c). The magnitude of HFA also differed among the
three tree species (Table 1), being greatest for pine (Fig. 2d),
intermediate for aspen (Fig. 2e) and least for spruce (Fig. 2f).
Pine, aspen and spruce all had periods where respiration was
signicantly greater at home. However, both aspen (day 954)
and spruce (day 61127) also had periods where there was a
negative additional respiration at home (i.e. home-eld disad-
vantage), and by day 225, there was no additional cumulative
respiration at home for these species (i.e. values did not differ
signicantly from zero).
FIELD EXPERIMENT
During winter, when the forest oor was snow-covered, soil
temperature was similar in stands of all three tree species at all
times of the day (Fig. 3). However, following snowmelt, day-
time temperatures were highest in aspen stands and lowest in
spruce stands, particularly during spring and summer
(P < 0.001 for time of day month species interaction;
Fig. 3). There were also differences in soil moisture among
stands of the three tree species, with soil moisture being great-
est in spruce stands and least in pine stands (P < 0.001;
Fig. 3).

 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
906 E. Ayres et al.

(a) (c) (e)

1.5

(mmol CO2 microcosm1)

Additional resp. at home
Cumulative resp. (mmol 15 * * * * ** * * *
CO2 microcosm1) 1.0

10 0.5

0.0
5
0.5

0 1.0
(b) (d) (f)

(mmol CO2 microcosm1)

Additional resp. at home
1.5
Cumulative resp. (mmol

15 * * * * * * * *
CO2 microcosm1)

1.0
10 0.5

0.0
5
0.5
0 1.0
0 100 200 0 100 200 0 100 200
Day Day Day

Fig. 2. Cumulative respiration from microcosms containing (a) each litter type regardless of soil community and (b) each soil community regard-
less of litter type (aspen: solid line, pine: dashed line, spruce: dotted line; soil only: grey line) and additional respiration at home for (c) all litter
types combined, (d) pine litter, (e) aspen litter and (f) spruce litter. In c-f, positive values correspond to greater than expected respiration at home
(i.e. home-eld advantage), while negative values correspond to lower than expected respiration at home, and asterisks indicate signicant differ-
ences from zero (P < 0.05) across the ve sampling dates beneath them (denoted by the horizontal bars). Values are mean SE.

Mass loss was greatest from aspen litter (43% mass loss after
2 years) and least from spruce litter (27% mass loss after
2 years; Table 2, Fig. 4a) and greatest in aspen and spruce
stands and least in pine stands (Table 2, Fig. 4b). In addition,
there was a signicant interaction between litter type and stand
indicating the potential for greater than expected mass loss
from litter at home (Table 2). Indeed, there was signicantly
greater mass loss from all litter types combined when litter was
incubated in its home stand over the ve sampling dates
(Fig. 4c). Consistent with results from the laboratory experi-
ment, an additional mass loss at home was greatest for pine
(Fig. 4df).
Aspen, pine and spruce litter lost 16, 59 and 61%, respec-
tively, of its initial N by the rst sampling date, but subse-
quently immobilized N resulting in a net increase in litter N
after 2 years (Table 2, Fig. 5a). At the nal sampling date, N
immobilization was greatest in spruce litter and least in pine lit-
ter, and was greatest in pine stands and least in aspen stands
(Table 2, Fig. 5a). Similar to litter mass loss, there was a signi-
cant interaction between litter type and stand in relation to N
loss (Table 2). In this case signicantly less N was lost (or more
was immobilized) from pine litter at home, while additional N
loss at home did not differ from zero for aspen or spruce litter
(Fig. 5cf).
In situ below-ground respiration rates varied by stand, but
not by litter type (Table 2, Fig. 6a, b). Respiration rates were
signicantly higher in aspen stands than in stands of pine and
spruce. As with litter mass loss, there was a signicant interac-
tion between litter type and stand, potentially indicating
greater respiration rates when litter was incubated in its home
stand (Table 2). Mean additional respiration at home values
was positive at each sampling date for all litter types combined,
but only approached being signicantly greater than zero
(P = 0.09). However, an additional respiration at home was
signicantly greater than zero for pine (Fig. 6d), but not for
aspen or spruce (Fig. 6e, f).
Ammonium was the dominant form of inorganic N,
accounting for 91% of soil inorganic N (data not shown).
Soil inorganic N concentrations were greatest in spruce
stands and lowest in aspen stands, but did not differ with lit-
ter type (Table 2, Fig. 7a, b). Additional soil inorganic N at
home was not signicantly different from zero for all litter
types combined, pine or aspen (Fig. 7ce), but was signi-
cantly greater than zero for spruce (Fig. 7f). See Figs S2, S3,
S4 and S5 for graphs of mass loss, N loss, respiration and
soil inorganic N concentration for each litter type in stands
of each tree species.
Discussion
Several authors have speculated that soil biota associated with
particular plant species specialize in decomposing litter pro-
duced by that plant species (Bocock et al. 1960; Wardle 2002;
Ayres et al. 2009a) and our laboratory experiment, and two
other recent studies (Strickland et al. 2009a,b), validate this
hypothesis. When all litter types were considered together, the
additional respiration at home accounted for 2.2% of total
CO2 production from litter incubated with its home soil biota.
However, there was a considerable difference in the magnitude
of the HFA among the three species. After 225 days, the addi-
tional respiration at home for pine litter accounted for 7% of
total CO2 production in microcosms containing litter and soil
biota from pine stands, while there was no signicant HFA for
aspen and spruce at this stage. This pattern was mirrored in the

 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
 Home-eld advantage hastens decomposition 907

(a) 20 Our data also suggests that the relationship between HFA
and time is more complex than the simple pattern we predicted
16
Soil temperature (C)
in Fig. 1. For example, both aspen and spruce litter, but not
12 pine, exhibited an HFA at the beginning of the laboratory
experiment. One possible explanation for this is that soil biota
8
from aspen and spruce stands have developed characteristics
4 that allow them to rapidly colonize (and start decomposing)
0
aspen and spruce litter, respectively. Pine litter did not show
this early peak in HFA, possibly because lodgepole pine was
4 introduced to this region recently (Paulson & Baker 2006), and
as a result there might not have been sufcient time for the soil
(b) 20
community to develop characteristics to colonize it rapidly.
16 However, this explanation is inconsistent with the large HFA
Soil temperature (C)

that was ultimately observed for pine litter over the duration of
12
the laboratory experiment. A second unexpected observation
8 in the relationship between HFA and time, was the negative
4
HFA (or home-eld disadvantage) observed on some occa-
sions for aspen and spruce litter. Although it is not clear what
0 caused the negative HFA, one possibility is that changes in lit-
4
ter structure and chemistry during decomposition result in the
litter becoming more similar to another litter type at some
(c) 120 stages, which would allow an away soil community to decom-
pose it rapidly.
Soil moisture (%)

90 Our data provide some support for the hypothesis that

60
30
0 dance and composition of other important compounds (e.g.
0 400 800
Day
Fig. 3. Mean monthly soil temperature at (a) 3am and (b) 3pm and (c)
gravimetric soil moisture content at each sampling date in stands of
aspen (solid line), pine (dashed line) and spruce (dotted line). Values
are mean SE.
eld experiment, where only pine litter had signicantly greater
mass loss and respiration rates at home. For example, after
2 years of decomposition in the eld, 16% of total pine litter
mass loss at home could be accounted for by HFA. Therefore,
it is apparent that HFA can have a sizable effect on the decom-
position of some litter types.
We speculated that HFA would initially increase, plateau
and nally decrease as decomposition proceeded (Fig. 1b).
This pattern was supported by the HFA for pine litter in the
laboratory experiment, which increased over the course of the
experiment, but appeared to be approaching a plateau by day
225. Similarly, although the HFA for aspen litter never
reached the magnitude that was observed for pine litter, aspen
HFA peaked between days 119 and 176, and subsequently
declined. However, spruce litter did not exhibit this relation-
ship. It is difcult to investigate the relationship between HFA
and time in the eld experiment due to the limited number of
sampling dates and reduced experimental control; nonetheless,
the patterns do not seem inconsistent with those observed in
the laboratory experiment.
HFA increases with decreasing litter quality, since litter quality
(based on lignin:N) was lowest for pine and HFA was greatest
for this species. However, HFA was greater for aspen than for
spruce, despite the lower quality of spruce litter. A more com-
prehensive assessment of litter quality, including the abun-
phenolics), might be necessary to accurately predict HFA for
different litter types. Strickland et al. (2009a,b) also observed
an increase in HFA with decreased litter quality, with HFA
being largest for recalcitrant Rhododendron maximum litter,
intermediate for moderately recalcitrant Pinus taeda litter, and
smallest for relatively labile grass litter (Hordeum murinum or
Panicum virgatum). The fact that Strickland et al. (2009a,b)
observed a clearer relationship between litter quality and the
magnitude of HFA than we found in our study probably
results from the greater dissimilarity in the litter types they used
(i.e. tree and grass litter rather than just tree litter). In addition,
other factors may also contribute to the magnitude of HFA,
such as the degree of dissimilarity in soil community composi-
tion (or functional traits) among the sites, since it seems unli-
kely that highly dissimilar communities will be equally efcient
at decomposing a particular litter type.
As with the mass loss and respiration HFA, the N loss HFA
was most pronounced for pine litter. However, rather than
accelerating N loss, pine litter had lower N loss (or greater
immobilization) than expected when at home, which was con-
sistent with our hypothesis of changes in litter N during the
early stages of decomposition (Fig. 1d). However, if pine litter
mass loss occurs more rapidly at home than away throughout
the decomposition process, we expect that litter N loss will be
greater at home than away after a longer period of time, e.g.
after 5 years (Fig. 1d). Our ndings are in agreement with the
study by Hunt et al. (1988), who also observed greater litter N

 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
908 E. Ayres et al.

Table 2. P values from anovas of three measures of decomposition (litter mass loss, litter N dynamics and soil respiration) and soil inorganic N,
and the HFA corresponding to those measures (ADH in eqn 1), from the eld experiment. The stand treatment does not apply to the additional
decomposition at home or additional inorganic N at home values. Note that P values relating to additional mass loss, N loss, respiration or soil
inorganic N at home refer to differences among treatments, not whether there is a signicant HFA. Asterisks on the corresponding graphs
(Figs 4, 5, 6, 7) indicate whether there is a signicant HFA (i.e. whether the data is signicantly different from zero)

Additional Additional Additional Soil Additional soil

Litter litter mass Litter litter N loss respiration inorganic inorganic N
mass loss loss at home N loss at home Respiration at home N at home

Litter <0.001 0.596 <0.001 0.041 0.950 0.060 0.111 0.074

Stand <0.001 <0.001 0.001 0.027
Day <0.001 0.462 <0.001 0.580 <0.001 0.598 <0.001 0.873
Litter stand <0.001 <0.001 0.002 0.316
Litter day 0.013 0.398 <0.001 0.098 0.397 0.038 0.573 0.574
Stand day 0.962 0.006 0.172 0.081
Litter stand day 0.449 0.038 0.133 0.983
Block <0.001 0.556 <0.001 0.377 <0.001 0.734 0.219 0.976
Stand block <0.001 <0.001 0.003 0.215
Litter block <0.001 0.014 <0.001 <0.001 0.136 0.002 0.067 0.795
Day block 0.180 0.718 0.002 0.981 0.001 0.647 0.004 0.979
Litter stand block 0.011 <0.001 0.006 0.952
Litter day block 0.685 0.038 0.989 0.981
Stand day block 0.737 0.151 0.095 0.002

(a) (c) (e)

100
Additional mass loss
Mass remaining (%)

at home (% initial)

90 8 *
80 4
70
0
60
50 4
(b) (d) (f)
100
Mass remaining (%)

Additional mass loss

at home (% initial)

90 8 *
80 4
70
0
60
50 4
0 400 800 0 400 800 0 400 800
Day Day Day

Fig. 4. Litter mass remaining for (a) each litter type regardless of stand and (b) each stand regardless of litter type (aspen: solid line, pine: dashed
line and spruce: dotted line) and additional litter mass loss at home for (c) all litter types combined, (d) pine litter, (e) aspen litter and (f) spruce lit-
ter. In cf, positive values correspond to greater than expected mass loss at home (i.e. home-eld advantage), while negative values correspond to
lower than expected mass loss at home, and asterisks indicate signicant differences from zero (P < 0.05) across all sampling dates. Values are
mean SE.

immobilization at home than away during the early stages of
decomposition. Interestingly, although decomposing at home
inuenced litter N loss for pine, it did not affect soil inorganic
N concentrations beneath the litter. However, soil inorganic N
concentrations were greater than expected at home for spruce
litter, despite the fact that spruce exhibited minimal HFA in
relation to the other measures. It is not clear what caused this
response.
The magnitude of the mass loss HFA that we identied for
aspen, pine and spruce falls within the range of values calcu-
lated in a literature study of HFA in forest ecosystems, where
mass loss ranged from 9% slower to 29% faster at home than
away; mean: 8% faster at home (Ayres et al. 2009a). Thus,
while it is clear that the primary factors that control decomposi-
tion rates are climate and litter quality (Aerts 1997; Gholz et al.
2000; Trofymow et al. 2002; Parton et al. 2007; Cornwell et al.

 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
 Home-eld advantage hastens decomposition 909

(a) (c) 20 (e)

Additional N loss at
125

home (% initial)
N remaining (%)
*
100 0
75
50 20
25
0 40

(b) (d) 20 (f)

Additional N loss at
125 *

home (% initial)
N remaining (%)

100 0
75
50 20
25
0 40
0 400 800 0 400 800 0 400 800
Day Day Day

Fig. 5. Litter nitrogen remaining for (a) each litter type regardless of stand and (b) each stand regardless of litter type, and additional litter nitro-
gen loss at home for (c) all litter types combined, (d) pine litter, (e) aspen litter and (f) spruce litter. In cf, positive values correspond to greater
than expected N loss (or lower than expected N immobilization) at home, while negative values correspond to lower than expected N loss (or
greater than expected N immobilization) at home, and asterisks indicate signicant differences from zero (P < 0.05) across all sampling dates.
Values are mean SE.

(a) (c) (e)

Additional respiration at

0.8
home (g CO2 m2 h1)

1.0
(g CO2 m2 h1)
Respiration rate

0.8
0.4
0.6
0.4
0.0
0.2
0.0 0.4
(b) (d) (f)
0.8
1.0
Additional respiration at
home (g CO2 m2 h1)

*
(g CO2 m2 h1)
Respiration rate

0.8
0.4
0.6
0.4
0.0
0.2
0.0 0.4
0 400 800 0 400 800 0 400 800
Day Day Day

Fig. 6. Soil respiration rate for (a) each litter type regardless of stand and (b) each stand regardless of litter type (aspen: solid line, pine: dashed line
and spruce: dotted line) and additional respiration at home for (c) all litter types combined, (d) pine litter, (e) aspen litter and (f) spruce litter. In
cf, positive values correspond to greater than expected respiration at home (i.e. home-eld advantage), while negative values correspond to
lower than expected respiration at home, and asterisks indicate signicant differences from zero (P < 0.05) across all sampling dates. Soil respi-
ration was not measured at the last sampling date due to inclement weather. Values are mean SE.

2008), HFA should be recognized as a secondary factor
controlling decomposition. Indeed, the inuence of HFA on
decomposition is comparable in magnitude to the effect of
litter interactions in mixtures (McTiernan, Ineson & Coward
1997; Wardle, Bonner & Nicholson 1997; Gartner & Car-
don 2004). For example, in a review of studies involving lit-
ter decomposing alone or in mixtures, Gartner & Cardon
(2004) reported that of 162 litter mixtures, mass loss was
slower than expected in 31 cases (on average 9% slower),
faster than expected in 77 cases (on average 17% faster),
while mass loss occurred at the expected rates in 54 cases.
In addition to demonstrate the role that HFA plays in litter
decomposition, our data also challenge two widely held views
in soil ecology: (i) soil biota are largely functionally redundant;

 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
910 E. Ayres et al.
(a)
Soil inorganic N (g g1)
15
10
5
0 6
(b)
15
Soil inorganic N (g g1)
10
5
0
0 400
Day
Fig. 7. Soil inorganic N concentration beneath litterbags for (a) each litter typ e regardless of stand and (b) each stand regardless of litter type
(aspen: solid line, pine: dashed lin e and spruce: dotted line) and additional soil inorganic N at home for (c) all litter types combined, (d) pine litter,
(e) aspen litter and (f) spruce litter. In cf, positive values correspond to greater than expected inorganic N at home, while negative values corre-
spond to lower than expected inorganic N at home and asterisks indicate signicant differences from zero (P < 0.05) across all sampling dates.
Values are mean SE.
and (ii) decomposition is a highly non-specic process. A few
handfuls of soil typically contain thousands of microbial spe-
cies and hundreds of animal species (Torsvik, Goksyr & Daae
1990; Bloemers et al. 1997; Gans, Wolinsky & Dunbar 2005; St
John, Wall & Hunt 2006; Wu et al. 2009) and several authors
have concluded that most of these soil organisms are function-
ally redundant (Andren, Bengtsson & Clarholm 1995; Groff-
man & Bohlen 1999; Setala, Berg & Jones 2005). This is
because experimental manipulations of soil species richness
have revealed only very weak effects on ecosystem functions
once 10 or more species are present in the community (Grifths
et al. 2000; Liiri et al. 2002; Setala & McLean 2004). However,
our study indicates that soil communities associated with dif-
ferent plant species are not functional equivalent, which is why
we observed HFA. Strickland et al. (2009a) reached the same
conclusion based on similar results from their reciprocal trans-
plant experiment. Thus, while functional redundancy may be
high for species within a community (Grifths et al. 2000; Liiri
et al. 2002; Setala & McLean 2004), functional redundancy
may be lower among species that are present in different soil
communities (e.g. communities associated with different plant
species). Moreover, we observed functional differences among
these soil communities in relation to decomposition, which is
generally considered to be a highly non-specic process and, as
a result, relatively insensitive to changes in soil community
composition (Grifths et al. 2000).
As well as providing information on HFA during decompo-
sition, our reciprocal litter transplant experiments also
revealed that both the site and the litter identity inuence
decomposition, which is similar to many other studies (Mc-
Claugherty et al. 1985; Hunt et al. 1988; Aerts 1997; Wardle,
Bonner & Nicholson 1997; Gholz et al. 2000; Trofymow et al.
2002; Hobbie et al. 2006; Parton et al. 2007; Cornwell et al.
 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
(c) (e)
18
Additional soil inorganic
N at home g g1)
12
6
0
(d) (f)
18
Additional soil inorganic
N at home g g1)
12
*
6
0
800 6
0 400 800 0 400 800
Day
Day
2008). Aspen litter decomposed faster than either pine or
spruce litter, a result that matches our litter quality data and is
in agreement with a previous study of the decomposition of
these litter types (Stump & Binkley 1993) and of litter from
related species (McClaugherty et al. 1985). Decomposition also
occurred more rapidly in stands of aspen and spruce, than in
pine stands. In part, this may have been caused by greater soil
temperature in aspen stands and greater soil moisture in spruce
stands, both of which are known to inuence decomposition
rates (Russell & Voroney 1998; Chapin, Matson & Mooney
2002; Hobbie et al. 2006). We also observed differences in N
immobilization in litter decomposing in the different stands,
similar to other multisite decomposition experiments (Hunt
et al. 1988; Hobbie & Vitousek 2000). This may have partly
resulted from differences in the rate mass loss among stands of
the three tree species, since mass loss is related to N immobili-
zation (Parton et al. 2007). In addition, the laboratory experi-
ment indicated that biota from aspen and pine stands
decomposed litter more rapidly than spruce soil biota. While
the decomposition capacity of these soil communities has not
been investigated previously, other studies have shown that
soil communities associated with three European tree species
(Ayres, Dromph & Bardgett 2006) and three geographically
isolated North American ecosystems (Strickland et al. 2009a)
differ in their capacity to decompose leaf litter. This presum-
ably results from differences in the composition of soil commu-
nities associated with different plant species, since soil
community composition is known to vary among plant species
(Grifths et al. 1992; Grayston et al. 1998; Priha, Hallantie &
Smolander 1999; Porazinska et al. 2003; Bardgett & Walker
2004) and can inuence decomposition rates (Cragg & Bardg-
ett 2001; Grifths et al. 2001; Heemsbergen et al. 2004; Setala
& McLean 2004; Strickland et al. 2009a).

Here, we provide the rst detailed study of HFA in relation
to decomposition. We demonstrate that HFA is caused by dif-
ferences in the soil community associated with different plant
species. We also found support for the notion that HFA has a
humpback relationship over time, although it is apparent that
other factors inuence this relationship given the initially large
HFA observed in the laboratory experiment, as well as periods
of home-eld disadvantage. In addition to effects on mass loss
and respiration, we show that HFA also inuences N cycling
by increasing N immobilization during the early stages of
decomposition. Our data suggest that soil biota are not func-
tionally redundant at the community level, which raises the
possibility that HFA effects on decomposition could be dis-
rupted by land use change, species migration in response to cli-
mate change, and species introductions, if this results in plant
species being separated from the soil community they have tra-
ditionally been associated with.
Acknowledgements
Koren Nydick and other staff at the Mountain Studies Institute, Silverton,
Colorado, USA, greatly assisted with logistics associated with this study. Chris
Landry and Jason Neff provided advice on site selection. Justin Anderson,
Mike Brown, Chrissy Cook, Ali Morse, Breana Simmons, Lisa Smith, Mat-
thew Territ and Matthew Wallenstein helped with eld work. This study was
supported by a British Ecological Society Early Career Project Grant to E.A.
and H.S. and an US National Science Foundation Ecosystem Science award to
E.A., H.S., and Matthew Wallenstein.
References
Aerts, R. (1997) Climate, leaf litter chemistry and leaf litter decomposi-
tion in terrestrial ecosystems: A triangular relationship. Oikos, 79, 439
449.
Andren, O., Bengtsson, J. & Clarholm, M. (1995) Biodiversity and species
redundancy among litter decomposers. The Signicance and Regulation of
Soil Biodiversity (ed. H.P. Collins). pp. 141151. Kluwer Academic Pub-
lisher, Dordrecht.
Austin, A.T. & Vivanco, L. (2006) Plant litter decomposition in a semi-arid eco-
system controlled by photodegradation. Nature, 442, 555558.
Ayres, E., Dromph, K.M. & Bardgett, R.D. (2006) Do plant species encourage
soil biota that specialise in the rapid decomposition of their litter? Soil Biol-
ogy & Biochemistry, 38, 183186.
Ayres, E., Heath, J., Possell, M., Black, H.I.J., Kerstiens, G. & Bardgett, R.D.
(2004) Tree physiological responses to above-ground herbivory directly
modify below-ground processes of soil carbon and nitrogen cycling. Ecology
Letters, 7, 469479.
Ayres, E., Steltzer, H., Simmons, B.L., Simpson, R.T., Steinweg, J.M., Wallen-
stein, M.D. et al. (2009a) Home-eld advantage accelerates leaf litter decom-
position in forests. Soil Biology & Biochemistry, 41, 606610.
Ayres, E., Steltzer, H., Berg, S., Wallenstein, M.D., Simmons, B.L. & Wall,
D.H. (2009b) Tree species traits inuence soil physical, chemical, and biolog-
ical properties in high elevation forests. PLoS ONE, 4, e5964.
Bardgett, R.D. & Walker, L.R. (2004) Impact of coloniser plant species on the
development of decomposer microbial communities following deglaciation.
Soil Biology and Biochemistry, 36, 555559.
Bloemers, G.F., Hodda, M., Lambshead, P.J.D., Lawton, J.H. & Wanless,
F.R. (1997) The effects of forest disturbance on diversity of tropical soil nem-
atodes. Oecologia, 111, 575582.
Bocock, K.L., Gilbert, O., Capstick, C.K., Twinn, D.C., Waid, J.S. & Wood-
man, M.J. (1960) Changes in leaf litter when placed on the surface of soils
with contrasting humus types: 1) Losses in dry weight of oak and ash leaf lit-
ter. Journal of Soil Science, 11, 19.
Cebrian, J. (1999) Patterns in the fate of production in plant communities.
American Naturalist, 154, 449468.
Chapin, F.S., Matson, P.A. & Mooney, H.A. (2002) Principles of Terrestrial
Ecosystem Ecology. Springer-Verlag, New York.
 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
Home-eld advantage hastens decomposition 911
Chapman, S.K. & Koch, G.W. (2007) What type of diversity yields synergy
during mixed litter decomposition in a natural forest ecosystem? Plant and
Soil, 299, 153162.
Clarke, S.R. & Norman, J.M. (1995) Home ground advantage of individual
clubs in English soccer. Statistician, 44, 509521.
Coleman, D.C., Blair, J.M., Elliot, E.T. & Wall, D.H. (1999) Soil invertebrates.
Standard Soil Methods for Long-Term Ecological Research (eds G.P. Robert-
son, D.C. Coleman, C.S. Bledsoe & P. Sollins). pp. 349377. Oxford Univer-
sity Press, Oxford.
Cornwell, W.K., Cornelissen, J.H.C., Amatangelo, K., Dorrepaal, E., Eviner,
V.T., Godoy, O. et al. (2008) Plant species traits are the predominant control
on litter decomposition rates within biomes worldwide. Ecology Letters, 11,
10651071.
Cragg, R.G. & Bardgett, R.D. (2001) How changes in soil faunal diversity and
compostion within a trophic group inuence decomposition processes. Soil
Biology & Biochemistry, 33, 20732081.
Dijkstra, F.A. & Cheng, W.X. (2007) Interactions between soil and tree roots
accelerate long-term soil carbon decomposition. Ecology Letters, 10, 1046
1053.
Gans, J., Wolinsky, M. & Dunbar, J. (2005) Computational improvements
reveal great bacterial diversity and high metal toxicity in soil. Science, 309,
13871390.
Gartner, T.B. & Cardon, Z.G. (2004) Decomposition dynamics in mixed-spe-
cies leaf litter. Oikos, 104, 230246.
Gholz, H.L., Wedin, D.A., Smitherma, S.M., Harmon, M.E. & Parton, W.J.
(2000) Long-term dynamics of pine and hardwood litter in contrasting envi-
ronments: toward a global model of decomposition. Global Change Biology,
6, 751765.
Grayston, S.J., Wang, S.Q., Campbell, C.D. & Edwards, A.C. (1998) Selective
inuence of plant species on microbial diversity in the rhizosphere. Soil Biol-
ogy and Biochemistry, 30, 369378.
Grifths, B.S., Welschen, R., Vanarendonk, J.J.C.M. & Lambers, H. (1992)
The effect of nitrate-nitrogen supply on bacteria and bacterial-feeding fauna
in the rhizosphere of different grass species. Oecologia, 91, 253259.
Grifths, B.S., Ritz, K., Bardgett, R.D., Cook, R., Christensen, S., Ekelund, F.
et al. (2000) Ecosystem response of pasture soil communities to fumigation-
induced microbial diversity reductions: an examination of the biodiversity-
ecosystem function relationship. Oikos, 113, 279294.
Grifths, B.S., Ritz, K., Wheatley, R., Kuan, H.L., Boag, B., Christensen, S.
et al. (2001) An examination of the biodiversity-ecosystem function relation-
ship in arable soil microbial communities. Soil Biology and Biochemistry, 33,
17131722.
Groffman, P.M. & Bohlen, P.J. (1999) Soil and sediment biodiversity - Cross-
system comparisons and large-scale effects. BioScience, 49, 139148.
Hattenschwiler, S. & Gasser, P. (2005) Soil animals alter plant litter diversity
effects on decomposition. Proceedings of the National Academy of Sciences,
102, 15191524.
Heath, J., Ayres, E., Possell, M., Bardgett, R.D., Black, H.I.J., Grant, H. et al.
(2005) Rising atmospheric CO2 reduces sequestration of root-derived soil
carbon. Science, 309, 17111713.
Heemsbergen, D.A., Berg, M.P., Loreau, M., van Haj, J.R., Faber, J.H. &
Verhoef, H.A. (2004) Biodiversity effects on soil processes explained by
interspecic functional dissimilarity. Science, 306, 10191020.
Hobbie, S.E. & Vitousek, P.M. (2000) Nutrient limitation of decomposition in
Hawaiian forests. Ecology, 81, 18671877.
Hobbie, S.E., Reich, P.B., Oleksyn, J., Ogdahl, M., Zytkowiak, R., Hale, C.
et al. (2006) Tree species effects on decomposition and forest oor dynamics
in a common garden. Ecology, 87, 22882297.
Hunt, H.W., Ingham, E.R., Coleman, D.C., Elliot, E.T. & Reid, C.P.P. (1988)
Nitrogen limitation of production and decomposition in prairie, mountain
meadow, and pine forest. Ecology, 69, 10091016.
Jamieson, D.W., Romme, W.H. & Somers, P. (1996) Biotic communities of the
cool mountains. The Western San Juan Mountains: Their Geology, Ecology,
and Human History (eds R. Blair, T.A. Casey, W.H. Romme & R.N. Ellis).
pp. 159174. University Press of Colorado, Niwot.
Laakso, J. & Setala, H. (1999) Sensitivity of primary production to changes in
the architecture of belowground food webs. Oikos, 87, 5764.
Liiri, M., Setala, H., Haimi, J., Pennanen, T. & Fritze, H. (2002) Relationship
between soil microarthropod species diversity and plant growth does not
change when the system is disturbed. Oikos, 96, 137149.
McClaugherty, C.A., Pastor, J., Aber, J.D. & Melillo, J.M. (1985) Forest litter
decomposition in relation to soil-nitrogen dynamics and litter quality. Ecol-
ogy, 66, 266275.
McTiernan, K.B., Ineson, P. & Coward, P.A. (1997) Respiration and nutrient
release from tree leaf litter mixtures. Oikos, 78, 527538.
912 E. Ayres et al.
Negrete-Yankelevich, S., Fragoso, C., Newton, A.C., Russell, G. & Heal, O.W.
(2008) Species-specic characteristics of trees can determine the litter macro-
invertebrate community and decomposition process below their canopies.
Plant and Soil, 307, 8397.
Parton, W.J., Silver, W.L., Burke, I.C., Grassens, L., Harman, M.E., Currie,
W.S. et al. (2007) Global-scale similarities in nitrogen release patterns during
long-term decomposition. Science, 315, 361364.
Paulson, D.D. & Baker, W.L. (2006) The Nature Of Southwestern Colorado.
University Press of Colorado, Boulder.
Porazinska, D.L., Bardgett, R.D., Blaauw, M.B., Hunt, H.W., Parsons, A.N.,
Seastedt, T.R. et al. (2003) Relationships at the aboveground-belowground
interface: Plants, soil biota, and soil processes. Ecological Monographs, 73,
377395.
Prescott, C.E., Zabek, L.M., Staley, C.L. & Kabzems, R. (2000) Decomposi-
tion of broadleaf and needle litter in forests of British Columbia: inuences
of litter type, forest type, and litter mixtures. Canadian Journal of Forest
Research, 30, 17421750.
Priha, O., Hallantie, T. & Smolander, A. (1999) Comparing microbial biomass,
denitrication enzyme activity, and numbers of nitriers in the rhizospheres
of Pinus sylvestris, Picea abies and Betula pendula seedlings by microscale
methods. Biology and Fertility of Soils, 30, 1419.
Russell, C.A. & Voroney, R.P. (1998) Carbon dioxide efux from the oor of a
boreal aspen forest. I. Relationship to environmental variables and estimates
of C respired. Canadian Journal of Soil Science, 78, 301310.
Ryan, M.G., Melillo, J.M. & Ricca, A. (1990) A comparison of methods for
determining proximate carbon fractions of forest litter. Canadian Journal of
Forest Research, 20, 166171.
Setala, H., Berg, M.P. & Jones, T.H. (2005) Trophic structure and functional
redundancy in soil communities. Biological Diversity and Function in Soils
(eds R.D. Bardgett, M.B. Usher & D.W. Hopkins). pp. 236249. Cambridge
University Press, Cambridge.
Setala, H. & McLean, M.A. (2004) Decomposition rate of organic substrates in
relation to the species diversity of soil saprophytic fungi. Oecologia, 139, 98
107.
St John, M.G., Wall, D.H. & Hunt, H.W. (2006) Are soil mite assemblages
structured by the identity of native and invasive alien grasses? Ecology, 87,
13141324.
Strickland, M.S., Lauber, C., Fierer, N. & Bradford, M.A. (2009a) Testing the
functional signicance of microbial community composition. Ecology, 90,
441451.
Strickland, M.S., Osburn, E., Lauber, C., Fierer, N. & Bradford, M.A. (2009b)
Litter quality is in the eye of the beholder: initial decomposition rates as a
function of inoculum characteristics. Functional Ecology, 23, 627636.
Stump, L.M. & Binkley, D. (1993) Relationships between litter quality and
nitrogen availability in Rocky Mountain forests. Canadian Journal of Forest
Research, 23, 492502.
Torsvik, V., Goksyr, J. & Daae, F.L. (1990) High diversity in DNA of soil
bacteria. Applied and Environmental Microbiology, 56, 782787.
 2009 The Authors. Journal compilation  2009 British Ecological Society, Journal of Ecology, 97, 901912
Trofymow, J.A., Moore, T.R., Titus, B., Prescott, C., Morrison, I., Siltanen,
M. et al. (2002) Rates of litter decomposition over 6 years in Canadian for-
ests: inuence of litter quality and climate. Canadian Journal of Forest
Research, 32, 789804.
Vivanco, L. & Austin, A.T. (2008) Tree species identity alters forest litter
decomposition through long-term plant and soil interactions in Patagonia,
Argentina. Journal of Ecology, 96, 727736.
Wall, D.H., Bradford, M.A., St. John, M.G., Trofymow, J.A., Behan-Pelletier,
V., Bignell, D.E. et al. (2008) Global decomposition experiment shows soil
animal impacts on decomposition are climate dependent. Global Change
Biology, 14, 26612677.
Wardle, D.A. (2002) Communities and Ecosystems: Linking the Aboveground
and Belowground Components. Princeton University Press, Princeton.
Wardle, D.A., Bonner, K.I. & Nicholson, K.S. (1997) Biodiversity and
plant litter: Experimental evidence which does not support the view
that enhanced species richness improves ecosystem function. Oikos, 79,
247258.
Wu, T.H., Ayres, E., Li, G., Bardgett, R.D., Wall, D.H. & Garey, J.R. (2009)
Molecular proling of soil animal diversity in natural ecosystems: Incongru-
ence of molecular and morphological results. Soil Biology & Biochemistry,
41, 849857.
Received 8 March 2009; accepted 17 June 2009
Handling Editor: Richard Bardgett
Supporting Information
Additional Supporting Information may be found in the online ver-
sion of this article:
Fig. S1. Cumulative respiration for each littersoil biota combination.
Fig. S2. Litter mass loss for each littersite combination.
Fig. S3. Litter nitrogen loss for each littersite combination.
Fig. S4. Belowground respiration for each littersite combination.
Fig. S5. Soil inorganic nitrogen concentrations beneath litterbags for
each littersite combination.
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