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Estimation of stand-level leaf area for boreal bryophytes

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Oecologia
DOI 10.1007/s00442-006-0619-5
E C O P H Y SI O L OG Y
Estimation of stand-level leaf area for boreal bryophytes
Ben Bond-Lamberty · Stith T. Gower
Received: 29 August 2006 / Accepted: 8 November 2006
© Springer-Verlag 2006
Abstract Bryophytes dominate the carbon and nitro-
gen cycling of many poorly drained terrestrial ecosys-
tems and are important in the vegetation-atmosphere
exchange of carbon and water, yet few studies have
estimated their leaf area at the stand scale. This study
quantiWed the bryophyte-speciWc leaf area (SLA) and
leaf area index (LAI) in a group of diVerent-aged
boreal forest stands in well and poorly drained soils.
Species-speciWc SLA (for three feather mosses, four
Sphagnum spp. and Aulacomnium palustre mixed with
Tomentypnum nitens) was assessed by determining the
projected area using a Xatbed scanner and cross-sec-
tional geometry using a dissecting microscope. The
hemisurface leaf area was computed as the product of
SLA and live biomass and was scaled by coverage data
collected at all stands. Pleurozium schreberi dominated
the spatial coverage, biomass and leaf area in the well-
drained stands, particularly the oldest, while S. fuscum
and A. palustre were important in the poorly drained
stands. Live moss biomass ranged from 47 to 230 g m¡2
in the well-drained stands dominated by feather
mosses and from 102 to 228 g m¡2 in the poorly drained
stands. Bryophyte SLA varied between 135 and
473 cm2 g¡1, for A. palustre and S. capillifolium, respec-
tively. SLA was strongly and signiWcantly aVected by
bryophyte species, but did not vary between stands; in
general, there was no signiWcant diVerence between the
Communicated by Jim Ehleringer.
B. Bond-Lamberty (&) · S. T. Gower
Department of Forest Ecology and Management,
University of Wisconsin, 1630 Linden Drive,
Madison, WI 53706, USA
e-mail: bpbond@wisc.edu
SLA of non-Sphagnum mosses. Bryophyte LAI
increased with stand age, peaking at 3.1 and 3.7 in the
well and poorly drained stands, respectively; this repre-
sented approximately 40% of the overstory LAI in the
well-drained stands and 100–1,000% in the poorly
drained stands, underscoring the important role bryo-
phytes play in the water and carbon budgets of these
boreal forests.
Introduction
Bryophytes dominate the carbon and nitrogen cycling
of many poorly drained terrestrial ecosystems (Turet-
sky 2003; Vitt et al. 2001), particularly in the boreal
region, but on a global scale as well (Longton 1992;
O’Neill 2000). Such ecosystems—peatlands and to a
lesser degree forested wetlands—cover at least
2.5 £ 108 ha globally (Gorham 1991; Lugo et al. 1990;
Matthews and Fung 1987), store approximately 455 Pg
C, mostly in peat, and sequester nearly a third of global
soil C (Gorham 1991). Much of the boreal forest is
poorly drained, with poorly drained areas exhibiting
high rates of primary production (Camill et al. 2001;
Vitt et al. 2001), slow decomposition rates and bryo-
phyte-dominated successional pathways (O’Neill 2000;
Turetsky 2003). Bryophytes are increasingly recog-
nized to play important roles in the biogeochemical
cycling of well-drained boreal forests as well (DeLuca
et al. 2002; O’Connell et al. 2003a) and inXuence the
vegetation-atmosphere exchange in many ecosystems
(LaXeur and Rouse 1988; Shimoyama et al. 2004;
Williams and Flanagan 1996).
Bryophytes are less well studied than vascular
plants, and few studies have quantiWed the bryophyte
123

leaf area index (LAI), the amount of leaf area per unit
ground area. These few studies have generally mea-
sured the leaf area of individual samples (Proctor 2000;
Rice and Schneider 2004; Williams and Flanagan 1998;
Zotz et al. 2000), although DeLucia et al. (2003) esti-
mated stand-level bryophyte LAI in a New Zealand
forest. In most vascular plant systems, LAI strongly
inXuences energy, water and carbon dioxide exchange
between terrestrial ecosystems and the atmosphere and
is tightly coupled with photosynthesis, litterfall, micro-
climate and productivity (Gower et al. 1999). It is an
important parameter in terrestrial biogeochemical
models and remote sensing (Chen et al. 1997), with
many models requiring it as an input, and allows
researchers to scale Xuxes from the leaf to ecosystem
level (Campbell and Norman 1998). Bryophytes diVer
in many respects from vascular plants—they are gener-
ally astomatal, poikilohydric and at least partially ecto-
hydric (carry external water)—but the bryophyte
photosynthetic apparatus itself is similar to that of a C3
vascular plant (Green and Lange 1994; Williams and
Flanagan 1998). Bryophyte LAI may be much more
tightly coupled with carbon than water exchange (Proc-
tor 2000), unlike in most vascular plants, but knowl-
edge of the bryophyte leaf area would be valuable for
studies at the plant, ecosystem and landscape levels.
Many methods have been used to estimate the leaf
area of large vascular plants (Bonan 1993; Chen et al.
1997); most bryophyte studies of leaf area have used
contact probes (Rice and Schneider 2004) or light
attenuation measurements (DeLucia et al. 2003). Leaf
area may also be calculated from leaf mass using spe-
ciWc leaf area (SLA), the ratio of the fresh foliage sur-
face area to the unit dry foliage mass. In vascular plants
SLA typically varies with tree species and environmen-
tal conditions, decreases with foliage age and higher
canopy position, and is correlated with the potential
growth rate (Lambers and Poorter 1992) and leaf lon-
gevity (Gower et al. 1993; Reich et al. 1992, 1997).
Many studies of bryophyte biomass have been per-
formed, and knowledge of bryophyte SLA could pro-
vide bryophyte leaf area estimates for many areas. We
know of no published measurements of bryophyte
SLA, however.
The goals of this study were to quantify bryophyte
spatial coverage, live biomass, SLA and thus LAI for a
group of boreal forest stands. This was conducted on a
species-speciWc level as bryophyte diversity and mor-
phology are highly variable, as in vascular plants (Shaw
and GoVinet 2000). Because structural parameters such
as SLA can change drastically with water status (Tuba
et al. 1996), bryophyte water content was also sampled
and reported.
123
Oecologia
Methods
Site descriptions
The study was conducted in four diVerent-aged (16, 41,
75 and 155 years in 2005) boreal forests west of
Thompson, Manitoba, Canada; the oldest stand (at
55°53⬘N, 98°20⬘W) is the BOREAS NSA tower site
(Halliwell and Apps 1997). Separate well and poorly
drained stands (about 50 m £ 50 m) were located in
each forest. The overstory in the two oldest stands was
primarily black spruce; in the younger two stands,
trembling aspen (Populus tremuloides Michx) and jack
pine (Pinus banksiana Lamb.) were also present
(Bond-Lamberty et al. 2002b). In well-drained stands
of central Manitoba, the black spruce canopy closure,
at 50–60 years, is associated with drastic thinning of the
understory and growth of thick feather mosses; in the
poorly drained stands, the canopy remains open, with
black spruce, Labrador tea (Ledium groenlandicum
Oeder), and Sphagnum spp. dominating biomass and
net primary production (Bond-Lamberty et al. 2004;
Gower et al. 1997; Wang et al. 2003). Mean annual
temperatures were ¡1 to ¡2°C. Select stand character-
istics are summarized in Table 1.
Field sampling protocol
In each stand, the spatial coverage of bryophyte spe-
cies was assessed by randomly placing a 0.25 m2 sam-
pling ring on the ground (n = 100) and visually
estimating the percent cover of each species in the ring.
Approximately 1% of a 50 m £ 50 m area, i.e., 25 m2,
was sampled in this way. We did not attempt to enu-
merate every single species present, as some bryophyte
species require microscopic analysis to distinguish, and
others are extremely rare. Instead, we focused on the
three to six species in each stand that dominated
(95%+) spatial coverage, and presumably biomass, leaf
area and net primary production.
Weft-forming and pleurocarpous feather mosses cat-
egorized were Schreber’s big red stem moss [Pleuroz-
ium schreberi (Brid.) Mitt.], knight’s plume moss
[Ptilium crista-castrensis (Hedw.) De Not.] and splen-
did feather moss [Hylocomium splendens (Hedw.)
Schimp. in B.S.G.]. In the lowlands, Sphagnum species
enumerated included S. fuscum (Schimp.) Klinggr., S.
riparium Ångstr. and S. warnstorWi Russ., S. angustifo-
lium (C. Jens. ex Russ.) C. Jens. in Tolf and S. capillifo-
lium (Ehrh.) Hedw. Also common in both well and
poorly drained stands was glow moss, Aulacomnium
palustre (Hedw.) Schwaegr. This bryophyte has a simi-
lar growth form to, and frequently grows intermixed
Oecologia
Table 1 Stand characteristics, by soil drainage and stand age (years), with standard errors in parentheses
Well drained
16 41 75
Tree diameter 0.0 (-) 3.0 (0.9) 4.7 (0.9)
Tree basal area 0.0 (-) 11.6 (3.9) 36.7 (4.0)
Overstory LAI 0.0 (-) 1.8 (0.7) 6.8 (0.7)
Vegetation C 0.5 (0.2) 2.1 (0.5) 6.5 (1.3)
Bryophyte NPP 143.7 (78.2) 8.3 (7.3) 27.3 (8.8)
Total NPP 365.9 (94.9) 406 (49) 358 (64)
Carbon content data (kg C m¡2 ) are from Wang et al. (2003), and net primary production (NPP, g C m¡2 year¡1 ) from Bond-Lam-
berty et al. (2004). Other data, including the mean tree diameter (cm) and basal area (m2 ha¡1 ) and overstory leaf area index (LAI)
were reported in Bond-Lamberty et al. (2002a, b)
with, the brown moss Tomentypnum nitens (Hedw.)
Loeske (Johnson et al. 1995); we did not attempt to
separate these two species. An ‘other’ category was
used for infrequently occurring bryophytes, in these
stands most often Polytrichum spp. The names and tax-
onomy given here follow the USDA plants database
(http://www.plants.usda.gov).
Five random samples, each 10 cm £ 10 cm and gen-
erally approximately 10 cm depth, were taken of each
major (>5% coverage) bryophyte species at the three
oldest stands. Sampling occurred over a 4-day period in
August 2005 during which there was no rain; meteoro-
logical records from Thompson, MB (40 km east) indi-
cated 21 mm of rain in the previous 7 days and 135 mm
in the previous 30 days (D. Bronson, University of
Wisconsin, unpublished data). Samples were stored in
airtight plastic bags, chilled and processed in the lab
within 24 h.
Laboratory protocol
Samples were sorted, weighed and scanned in the labo-
ratory. Each sample was separated into live and dead
tissue based on color, depth and decay. Treatment of
samples was species-speciWc:
H. splendens: This species was separated into live
and dead tissue, and the live tissue separated into
1st- and 2nd-year growth; Hylocomium is the only
bryophyte sampled with such a determinate growth
form (Fig. 1). Leaves and secondary costae were
separated from the main midrib.
A. palustre/T. nitens, P. schreberi, P. crista-castren-
sis: The top approximately 1.5 cm was treated as
live, following Williams and Flanagan (1998) and
Weld observations (Bond-Lamberty, pers. obs). For
P. schreberi, H. splendens and A. palustre, the main
midrib was separated from secondary costae and
scanned and weighed separately.
Poorly drained
155 16 41 75 155
7.9 (1.2) 0.0 (-) 2.3 (0.4) 4.7 (1.3) 3.5 (0.5)
42.2 (8.6) 0.0 (-) 2.2 (1.0) 20.3 (5.1) 16.6 (6.1)
5.3 (0.9) 0.0 (-) 0.3 (0.1) 3.0 (0.8) 1.1 (0.4)
8.3 (1.6) 0.6 (0.1) 2.0 (0.4) 3.7 (0.8) 3.4 (0.7)
12.0 (6.0) 36.8 (28.6) 146.8 (88.8) 89.8 (20.2) 24.0 (12.0)
211 (50) 239.0 (68.9) 401 (105) 311 (54) 261 (49)
S. fuscum: The very compact growth form of this
species allows little light to penetrate below the
capitulum, and thus the top 0.5 cm was treated as
the live (photosynthesizing) plant. The branches of
S. fuscum are densely interleaved in fascicles and
were separated before scanning (Fig. 1).
S. warnstorWi, S. riparium, S. capillifolium: These
species are less densely arranged than S. fuscum,
and the top approximately 2.5 cm were treated as
live. Branches were separated before scanning.
Bryophyte samples were scanned at 600 DPI using a
backlit Xatbed scanner (Canon LiDE, Canon USA,
Lake Success, NY). The resulting TIFF image Wles
were converted from grayscale to black and white
using a threshold luminosity value of 175 (on a 0–255
scale). This threshold value yielded pixel counts virtu-
ally identical to a subset of images in which pixels were
counted by hand under both low-shadow (y = 0.9642x,
R2 = 0.997, n = 10) and high-shadow (y = 0.7132x,
R2 = 0.991, n = 10) conditions (Bond-Lamberty,
unpublished data). Two separate regressions were used
because some samples exhibited pronounced shadow-
ing, caused by the small bryophyte leaves not contact-
ing the scanner bed cover, but instead projecting their
shadows upon it. Images were subjectively categorized
as “low-shadow” and “high-shadow,” and a hand count
was performed if there was any doubt. Approximately
13% of the samples (n = 19) were hand-counted, with
pixel counts on average 1% (§SD of 11%) below the
regression-based value. Projected leaf area (PLA) was
computed by dividing the Wnal image pixel count
(based on the appropriate regression, or the hand
count) by the scan resolution in pixels per unit area.
Leaf area index (LAI) was deWned as hemisurface
leaf area (HSLA), or one-half total leaf area per unit
ground area. The conversion of PLA to HSLA is
dependent on the cross-sectional leaf geometry (Bond-
Lamberty et al. 2003; Chen et al. 1997). HSLA for Xat-
123
 Oecologia

Fig. 1 Microscope images of

bryophyte samples. Top row,
left to right: Pleurozium schre-
beri at 7.5£; P. schreberi at
30£; Hylocomium splendens’
determinate growth form at
7.5£. Bottom row, left to right:
Sphagnum fuscum at 7.5£;
separated branches of S. fu-
scum at 7.5£; detail of S. fu-
scum branch at 64£. Scale
ruler (orange lines) is in mm

leaved species (the three feather mosses H. splendens,
P. schreberi and P. crista-castrensis, and A. palustre)
was assumed to be equal to PLA (i.e.,
HSLA:PLA = 1). A dissecting microscope was used to
examine Sphagnum branch cross-sections (Fig. 1).
Based on these images and the known morphology of
Sphagnum (SchoWeld 2001), a semicircular geometry
was used for all Sphagnum species (HSLA:PLA =
/2).
All samples were dried to a constant mass in a
forced-air oven at 70°C, weighed and the water content
computed on a dry-mass basis (fresh mass minus dry
mass, divided by dry mass). SpeciWc leaf area (SLA)
was computed as HSLA (cm2) divided by leaf mass (g).
Area-normalized biomass (g C m¡2) was computed
using the spatial survey and biomass data; for species
with non-photosynthesizing midribs (P. schreberi, A.
palustre and H. splendens), foliage and total biomass
(Bf and Bt) were calculated separately. The leaf area
index (LAI), deWned as HSLA per unit ground area,
was computed by multiplying foliage biomass by SLA
for each species. Site-speciWc SLA values were used
when possible; if a particular species was present at a
site, but its SLA was not sampled at that site, the mean
from other sites was used.
Statistical analyses were conducted using SAS ver-
sion 8.1 software (SAS Institute Inc. 2001). Post-hoc
tests for species and site diVerences were performed
using a Tukey-Kramer protected test in PROC
MIXED with a base  level of 0.05; unless otherwise
noted, all calculations and statistical analyses used
bryophyte sample as the experimental unit (n = 100)
and a signiWcance level of  = 0.05.
Results
Coverage, structure and water content
P. schreberi dominated bryophyte ground cover in the
two oldest well-drained stands (69–84%) and was an
important component in the two oldest poorly drained
stands (Table 2). P. schreberi spatial coverage was sig-
niWcantly related to overstory leaf area (y = ¡3.2 +
12.4x, R2 = 0.93; P = 0.0001; data not shown). A. palus-
tre was the dominant bryophyte in the younger stands;
Sphagnum species, particularly S. fuscum, were impor-
tant in the poorly drained stands (Table 2). Bryophyte
diversity was higher in the poorly drained stands than
in the well drained ones, and monotonically increased
with stand age in the poorly drained stands.
Three of the bryophyte species sampled (A. palustre,
H. splendens and P. schreberi ) had relatively tough,

123
Oecologia

Table 2 Spatial coverage of

Species Well drained Poorly drained
brophyte species in study sites
(%), by soil drainage and 16 41 75 155 16 41 75 155
stand age (years)
A. palustre 30.8 3.6 0.0 0.0 33.5 43.1 23.1 23.3
H. splendens 0.0 4.5 0.0 16.7 0.0 0.0 0.0 6.8
Results are based on visual P. schreberi 1.0 4.5 83.9 68.6 0.0 2.4 21.6 20.1
inspection of 0.25 m2 samples P. crista-castrensis 0.0 0.0 10.7 2.0 0.0 0.0 0.0 0.0
(n = 100). Bryophyte abbrevi- S. capillifolium 0.0 0.0 0.0 0.0 0.0 1.7 3.5 2.0
ations are based on the Wrst S. fuscum 0.0 0.0 0.0 0.4 0.0 12.9 5.6 3.4
two letters of genus and spe- S. riparium 0.0 0.0 0.0 0.3 0.0 2.6 9.0 5.9
cies names and are given in S. angustifolium 0.0 0.0 0.0 1.0 0.0 1.6 1.5 1.1
text. The Wnal line of the table S. warnstorWi 0.0 0.0 0.0 0.0 0.0 3.3 6.7 10.1
gives species diversity as mea-
Other 7.0 0.8 0.3 1.6 0.0 4.9 2.3 5.5
sured by the Shannon Index
Total coverage 38.8 13.4 94.9 90.6 33.5 72.4 73.2 78.1
(Legendre and Legendre
Diversity 0.59 1.25 0.37 0.77 0.00 1.34 1.73 1.86
1998)

non-photosynthesizing main midribs, and leaves and
midrib masses were determined separately. Leaf mass
as a percentage of total sample mass was 37 § 5%
(mean § standard deviation, n = 10) for A. palustre,
54 § 12% (n = 8) for H. splendens and 55 § 9%
(n = 18) for P. schreberi. These percentage did not
diVer signiWcantly between stands (F3,12 = 1.81;
P = 0.199; two stands for A. palustre and H. splendens,
four stands for P. schreberi). Leaves of A. palustre
comprised a signiWcantly smaller fraction of plant mass
than did those of H. splendens or P. schreberi
(F1,12 = 17.31; P = 0.001).
Water content in situ—deWned as bryophyte fresh
mass minus dry mass, divided by dry mass—did not
diVer between the live moss at the surface and dead tis-
sue underneath (F1,151 = 0.01; P = 0.925). Water con-
tent did diVer signiWcantly between species, even after
taking into account site-to-site variability
(F6,151 = 100.48; P < 0.001); from highest to lowest
water content, the species’ order in this regard was
S. riparium (2,066% § 520%, n = 23) > S. capillifolium >
S. warnstorWi > S. fuscum > A. palustre > P. crista-
castrensis > P. schreberi > H. splendens (400 § 92%,
n = 73). The well-drained 41-year-old stand had drier
bryophytes than the others (T151 = ¡2.09; P = 0.039);
otherwise, there was no diVerence between stands in
this regard.
Biomass, speciWc leaf area and leaf area index
Total live bryophyte biomass in the well-drained stands
ranged from 46.6 g m¡2 at the 41-year-old stand to
229.9 g m¡2 at the 155-year-old one (Fig. 2). Biomass in
the poorly drained stand was generally higher, from
102.1 g m¡2 in the youngest stand to 228.0 g m¡2 at the
oldest. The dominance of A. palustre, H. splendens and
P. schreberi meant that in most stands total live bio-
mass was equally comprised of leaf and non-leaf tissue.
SpeciWc leaf area of samples varied between 135.1
and 472.9 cm2 g¡1; these lowest and highest values were
for A. palustre and S. capillifolium, respectively (both
at the 41-year-old poorly drained stand; Table 3). SLA
was strongly aVected by bryophyte species across the
entire data set (F7,61 = 26.44; P < 0.0001), but not by
stand (F5,61 = 1.10; P = 0.374); the species-stand inter-
action was not signiWcant. For individual bryophyte
species, the only signiWcant diVerence in SLA among
stands was for A. palustre (135.1 versus 206.3 and
212.8 cm2 g¡1; F2,9 = 13.17; P = 0.004; Table 3). In gen-
eral, there was no signiWcant diVerence between the
SLA of non-Sphagnum mosses.
The midrib of P. schreberi had signiWcantly lower
SLA (diVerence of 91.5 cm2 g¡1; F1,17 = 23.72;
P = 0.0001) than did the leaves, as did the midrib of A.
palustre (diVerence of 61.4 cm2 g¡1; F1,11 = 24.56;
P = 0.0004). The midrib of H. splendens also had lower
SLA than its leaves (by 39.1 cm2 g¡1), but the diVer-
ence was not signiWcant (F1,7 = 2.67; P = 0.146). Sphag-
num species had much more variable, and signiWcantly
(F2,51 = 24.29; P < 0.0001) higher, SLA than did the
feather mosses (Table 3). Leaf age (current year versus
previous year) had no signiWcant eVect on SLA of H.
splendens (F1,8 = 0.01; P = 0.920).
Bryophyte leaf area index increased with stand age
in both the well and poorly drained stands, peaking at
3.1 and 3.7, respectively (Fig. 3). Bryophyte LAI com-
prised 28–58% of overstory LAI in the well-drained
stands, but in the poorly drained stands bryophyte LAI
exceeded overstory LAI (by 336–1,133%) in every
stand (Bond-Lamberty et al. 2002b). Bryophyte LAI
was dominated by P. schreberi in the well-drained
stands and Sphagnum spp. in the poorly drained
stands. Total shoot area index (SAI, i.e., leaf + nonleaf
area, hemisurface basis) was 2.0–5.0 in the well-drained
stands and 1.6–4.9 in the poorly drained stands (data
not shown; SAI increased with stand age in both cases).

123
 Oecologia

Fig. 2 Bryophyte biomass by stand age, for a well-drained and b poorly drained stands. “Nonleaf” refers to main midrib of P. schreberi,
H. splendens and A. palustre. Error bars show standard deviation of total biomass (n = 100 samples, each 0.25 m2)

Table 3 SpeciWc leaf area (cm2 g¡1, hemisurface basis) for bryophyte species, by soil drainage and stand age (years)
Species Well drained Poorly drained

41 75 155 41 75 155

A. palustre 135.1a (25.8) 206.3a (19.5) 212.8a (19.1)

H. splendens 145.0a (13.9) 157.6a (40.6) 152.5a (16.8)
P. schreberi 262.1a (43.3) 196.8a (35.1) 242.6a (77.4) 242.8ab (15.1) 196.9a (14.0)
P. crista-castrensis 223.6a (27.6)
S. capillifolium 472.9b (73.6) 454.3bc (50.8)
S. fuscum 247.8c (31.9) 290.4a (72.5)
S. riparium 357.6b (60.7)
S. warnstorWi 362.9b (65.6) 306.9ab (61.9)
Standard deviations, in parentheses, are generally based on n = 5 samples. Bryophyte abbreviations are based on Wrst two letters of ge-
nus and species; see text. Superscript letters indicate signiWcant diVerences between species at a given site (i.e., within a column; Tukey-
Kramer protected test with a base  level of 0.05)

Discussion tive of a larger group of forest stands sampled through-

Bryophytes have a large impact on the carbon, nitro-
gen and water cycling of northern ecosystems (O’Neill
2000; Turetsky 2003), with diVerent functional groups
(i.e., feather mosses and Sphagnum) having signiW-
cantly diVerent eVects (Bisbee et al. 2001). The spatial
coverage and moss biomass numbers reported in this
study are consistent with previous studies in these sites
(Gower et al. 1997; Wang et al. 2003) and other boreal
forests (Longton 1992; O’Connell et al. 2003b; O’Neill
2000). Bisbee et al. (2001) reported Sphagnum percent
cover was inversely correlated to overstory LAI for a
boreal Saskatchewan forest, much as the current study
found Pleurozium cover positively correlated to over-
story LAI. Although this study sampled only a small
area (2,500 m2) in each of a few boreal forest sites, the
stands studied here have been shown to be representa-
out north-central Manitoba (Bond-Lamberty et al.
2004).
Bryophyte canopy structure and leaf area
Bryophyte canopy structure strongly aVects water
uptake, storage and loss and varies considerably
among species (Rice et al. 2001). A key structural
metric is SLA; the SLA values reported here spanned a
wide range (135–473 g cm¡2), especially within the
Sphagnum samples (Table 3). Light and water exert
strong inXuences on Sphagnum canopy structure (Hay-
ward and Clymo 1983), and variation with a species can
be caused by, e.g,. ultraviolet radiation in S. fuscum
(Gehrke 1998). The values in Table 3 may represent
the low end of bryophyte SLA, particularly as we did
not measure at the scale of individual single-cell branch

123
Oecologia
Fig. 3 Bryophyte leaf area index (LAI, m2 m¡2), hemisurface ba-
sis, by stand age, for a well-drained and b poorly drained stands.
Line graphs show overstory LAI for each stand (Bond-Lamberty
leaves. Rice and Schneider (2004) used a contact probe
to measure SLA of approximately 1,700 cm2 g¡1 for the
cushion moss Leucobryum glaucum (S. Rice, personal
communication), an order of magnitude larger than the
values reported here.
The large contribution of bryophyte LAI to overall
stand LAI, and importance of bryophyte to boreal car-
bon (Harden et al. 1997) and water (Price et al. 1997)
Xuxes, emphasize the need to include water and carbon
Xuxes of bryophytes in many forest ecosystem analyses.
However, few direct measurements of bryophyte leaf
area have been made. DeLucia et al. (2003) estimated a
total LAI of 6 for bryophytes in a mature New Zealand
forest, based on light attenuation through the bryophyte
canopy. The LAI values reported here are expressed as
hemisurface leaf area, i.e., one-half of the total leaf area,
and thus our oldest stands’ values (3.1–3.7) are similar to
those measured by DeLucia et al. (2003). We are
unaware of other previous published estimates of stand-
level bryophyte leaf area, although values for individual
samples have been reported. Proctor (2000) and Proctor
and Tuba (2002) estimated LAI values of 6 for Tortula
(Syntrichia) intermedia, 18 for Mnium hornum and 20–
25 for Scleropodium (Pleurscleropodium) purum. Wil-
liams and Flanagan (1998) measured leaf area of P.
schreberi from a black spruce site in Saskatchewan.
They did not report these data, but imply that a total
shoot area index of 2.4 was typical (Williams and Flana-
gan 1998), overlapping our data from the 155-year-old
stand, samples from which had (hemisurface) LAI val-
ues ranging from 0.5 to 5.3 and SAI from 0.6 to 6.3.
The high LAI values noted above are characteristic
of bryophytes (Proctor 2000), the result of a diVerent
et al. 2002b). Error bars show standard deviation (n = 100 sam-
ples, each 0.25 m2)
set of ecophysiological tradeoVs than those faced by
large vascular plants. High LAI certainly maximizes
light interception for photosynthesis (Whitehead and
Gower 2001); it may also mitigate the slow (by a factor
of 10¡4 as compared to air) molecular diVusion of CO2
through these plants’ internally and externally carried
water volumes (Proctor 2000). Evaporation from bryo-
phyte leaves is controlled by diVusion through bound-
ary layers adjacent to plant surfaces, as in vascular
plants (Campbell and Norman 1998). However, (1) the
general absence of stomata mean that the boundary
layer is the major impediment to water loss (Proctor
1980) and (2) bryophyte cushion size and surface rough-
ness control boundary layer thickness—and thus water
relations and carbon gain (Zotz et al. 2000)—as the
small bryophyte leaves lie entirely within the cushion’s
laminar boundary layer at low to moderate wind speeds
(Rice and Schneider 2004). Thus, in most conditions the
bryophyte colony surface functions as a single “leaf”
lacking an outer epidermis (Proctor 2000), enabling the
bryophyte to maintain a large leaf area, while paying a
smaller penalty in water loss than would most vascular
plants. Their impact on ecosystem water cycling is
nonetheless signiWcant: boreal bryophytes were shown
to intercept 23% of water throughfall (compared to 15–
60% for the overstory canopy), with most of that subse-
quently evaporating, in a Picea mariana canopy near
the sites described here (Price et al. 1997).
Sources of error and methodology assessment
Several potential sources of error can be identiWed in our
measurements and estimates of stand-level bryophyte
123

LAI. First, the scale of measurement is a signiWcant
factor with bryophytes, whose functional units span
three orders of magnitude (0.1 to 10+ mm) (Proctor
and Tuba 2002; Rice et al. 2001). It is important to note
that our measurements were not at the scale of the
actual branch leaves in some of these species (Fig. 1),
aVecting this method’s accuracy. For example, the
actual branch leaves of Sphagnum are one cell thick
and overlap (SchoWeld 2001), and the actual hemisur-
face area of such cells can be expected to be larger (and
thus the SLA and LAI higher) than the values given
here. Higher PLA:HSLA conversion factors than the
ones used here, which assumed smooth surface geome-
tries, could be used to convert projected to hemisur-
face leaf area at this small scale.
A second problem is one of sampling design, not
methodology: as noted above, the results given for
A. palustre actually include a mix of this species and
T. nitens. These two mosses have some similarities
(and in this study there was no diVerence in non-
Sphagnum SLA), but belong to diVerent taxonomic
orders. Given the ubiquity of these species in boreal
fens and swamps (Johnson et al. 1995), and the great
variability in SLA measured here (Table 3), species-
speciWc data could be useful.
Thus, there are reasonable questions about the accu-
racy of our results. The precision of this method,
however, is high: biomass, projected area and cross-
sectional morphology can all be determined with rela-
tively little error. In addition, once SLA values are
determined, estimation of bryophyte LAI simply
requires an accurate biomass estimation—a simple and
routinely made measurement, and one that integrates
over the entire stand area. In contrast, techniques such
as contact probes (Rice and Schneider 2004) and light
extinction measurements (DeLucia et al. 2003) are sin-
gle-point measurements that require more time and
money to perform well. Direct comparisons of bryo-
phyte leaf area determined by these diVerent methods
would be a useful next step in accurately estimating
stand-level bryophyte LAI.
Finally, the ease of LAI estimation using this
method may in turn promote new research possibili-
ties. For instance, moss Xux measurements are typically
expressed on a ground area basis, unlike the leaf-level
measurements performed for vascular plants, but
knowledge of moss LAI would allow for more direct
comparisons to be made between the leaf gas exchange
rates of these groups. Currently, most studies of evapo-
transpiration do not consider bryophyte leaf area, even
when measuring moss-dominated systems (Shimoyama
et al. 2004). Mosses and other bryophytes play signiW-
cant roles in the biogeochemical cycling of boreal
123
Oecologia
systems, and a simple way to estimate their leaf area
would improve our understanding of these processes.
Acknowledgments This research was supported by grants from
NASA (NAG5-8069) and the National Science Foundation (Inte-
grated Research Challenges in Environmental Biology, DEB-
0077881) to S. T. Gower. We thank Frank Santiago, Denise Smith
and Glen Stanosz for their assistance in the Weld and lab and
appreciate the thoughtful comments of two anonymous reviewers.
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