Plant Ecology (2005) 177: 177–188
Springer 2005

DOI 10.1007/s11258-005-2322-8
-1

Biomass, production and woody detritus in an old coast redwood (Sequoia

sempervirens) forest

Richard T. Busing1,* and Takao Fujimori2

1
U.S. Geological Survey, 200 SW 35th St., Corvallis, OR 97333, USA; 2Japan Forest Technology Association, 7
Rokuban-cho, Chiyoda-ku, Tokyo, 102-0085, Japan; *Author for correspondence

Received 14 November 2003; accepted in revised form 08 June 2004

Key words: Decay, Natural disturbance, NPP, Old growth, Pacific Northwest, Stand dynamics, Woody debris

Abstract

We examined aboveground biomass dynamics, aboveground net primary production (ANPP), and woody detritus
input in an old Sequoia sempervirens stand over a three-decade period. Our estimates of aboveground biomass
ranged from 3300 to 5800 Mg ha
1. Stem biomass estimates ranged from 3000 to 5200 Mg ha
1. Stem biomass
declined 7% over the study interval. Biomass dynamics were patchy, with marked declines in recent tree-fall
patches <0.05 ha in size. Larger tree-fall patches approaching 0.2 ha in size were observed outside the study plot.
Our estimates of ANPP ranged from 6 to 14 Mg ha
1yr
1. Estimates of 7 to 10 Mg ha
1yr
1 were considered to
be relatively accurate. Thus, our estimates based on long-term data corroborated the findings of earlier short-term
studies. ANPP of old, pure stands of Sequoia was not above average for temperate forests. Even though
production was potentially high on a per stem basis, it was moderate at the stand level. We obtained values of 797
m3 ha
1 and 262 Mg ha
1 for coarse woody detritus volume and mass, respectively. Fine woody detritus volume
and mass were estimated at 16 m3 ha
1 and 5 Mg ha
1, respectively. Standing dead trees (or snags) comprised
7% of the total coarse detritus volume and 8% of the total mass. Coarse detritus input averaged 5.7 to 6.9 Mg
ha
1yr
1. Assuming steady-state input and pool of coarse detritus, we obtained a decay rate constant of 0.022 to
0.026. The old-growth stand of Sequoia studied had extremely high biomass, but ANPP was moderate and the
amount of woody detritus was not exceptionally large. Biomass accretion and loss were not rapid in this stand
partly because of the slow population dynamics and low canopy turnover rate of Sequoia at the old-growth stage.
Nomenclature: Hickman (1993).

Introduction ha
1yr
1. By comparison, mean ANPP of old tem-

Old Sequoia sempervirens forests have the greatest
terrestrial biomass levels on earth (> 3000 Mg ha
1;
Westman and Whittaker 1975; Fujimori 1977, 2001).
Whether their biomass dynamics are also distinctive is
unclear. The productivity of Sequoia forests appears
to resemble that of other old temperate forests
(Whittaker 1966). Westman and Whittaker’s (1975)
estimates of aboveground net primary production
(ANPP) in old Sequoia stands range from 5 to 19 Mg
perate forests is 10 to 13 Mg ha
1yr
1 (Whittaker
1966; Whittaker and Likens 1975). Even young
Sequoia stands tend to have unremarkable ANPP
(< 20 Mg ha
1yr
1; Westman and Whittaker 1975;
Olson et al. 1990). The few published estimates of
biomass and production thus far suggest that high
Sequoia forest biomass is not a consequence of high
productivity. It is the great longevity and size of
Sequoia trees that allow great biomass accumulation
(Whittaker 1966; Westman and Whittaker 1975).
178
Biomass loss and the dynamics of dead wood in
Sequoia forests have not been fully characterized
(Sawyer et al. 2000b). Bingham and Sawyer (1988)
noted large volume (957 m3 ha
1) and mass (200 Mg
ha
1) of coarse woody detritus in an old upland
Sequoia forest. Other estimates for Sequoia forests
range from 10 to 280 Mg ha
1 (Sawyer et al. 2000b).
However, rates of detritus input and decay have not
been widely reported. Sawyer et al. (2000b) speculate
that the decay rate of woody detritus is extremely slow
in Sequoia forests.
A three-decade study of the dynamics of compo-
sition and structure in an old Sequoia forest revealed
slow canopy turnover (Busing and Fujimori 2002).
Here we provide complementary information on
fundamental ecosystem processes. Our objectives are
to 1) estimate aboveground wood volume and bio-
mass, 2) examine the trajectory and spatiotemporal
dynamics of biomass, 3) estimate ANPP using several
methods, and 4) estimate woody detritus volume,
mass, input rate and decay rate. Implications of our
findings for the dynamics of old Sequoia ecosystems
are discussed. Furthermore, we compare and contrast
the fundamental ecosystem dynamics of our study
forest to those of other forests worldwide.
Study forest
Sequoia sempervirens occurs within 50 km of the Pa-
cific Ocean from southern Oregon to central
California. Particularly massive stands of Sequoia oc-
cur in the Central Redwood Forest, which ranges from
southern Humboldt County to San Francisco Bay
(Sawyer et al. 2000b). An old stand of very large trees
on alluvial flats within this biogeographic section was
selected for study. It lies near Bull Creek in Humboldt
Redwoods State Park (4021¢ N, 1230¢ W). Elevation
of the study site is 70 to 80 m above sea level. Climate,
geology and soils of the site are described by Fujimori
(1977). Moderate temperatures and high moisture in-
put are characteristic of the forest. Precipitation is high
except in summer. Fog drip compensates for low
summer precipitation (Byers 1953).
Sequoia comprises more than 99% of the total live
basal area in the study stand (Busing and Fujimori
2002). Overstory tree height often exceeds 90 m
(Fujimori 1977). Other tree species in the forest are
much shorter in stature. They include Umbellularia
californica, Lithocarpus densiflorus, Taxus brevifolia,
and Corylus cornuta.
Natural disturbances in the study forest include fire,
flooding and tree fall. Fire events are thought to cause
cumulative damage to Sequoia trees that may result in
death (Finney and Martin 1989). The presettlement
return interval for fire in Humboldt County and sur-
rounding areas is estimated at <10 to 500 yr (Veirs
1982; Stuart 1987). Veirs (1982) notes that mesic sites
near the coast experience light fires every 250 to 500
yr. Sites intermediate on the mesic-xeric continuum
have a fire return interval of 100 to 200 yr. Xeric
interior sites have a return interval of about 50 yr. The
study forest is likely to have a fire return interval
<100 yr (Stuart 1987). However, fire suppression
policies implemented in the 1900s have greatly re-
duced fire occurrence for at least the last 50 years
(Stone and Vasey 1968; Veirs 1982). By contrast,
flooding has occurred in the study stand in recent
decades (Sawyer et al. 2000a). The return interval for
floods covering most of the study stand is <100 yr.
Flooding replenishes nutrients (Westman and Whit-
taker 1975), deposits sediments, and influences
understory tree survival (Stone and Vasey 1968).
Establishment of Sequoia seedlings is common after
major floods (Sawyer et al. 2000a). Tree-fall gaps are
also an important form of disturbance in the study
forest. Although the return interval of gap disturbance
is at least 250 yr, individual gaps can exceed 0.1-ha in
area (Busing and Fujimori 2002). They currently
cover about 20% of the total area within the study
forest.
Methods
Biomass and production
In 1972, trees ‡10 cm diameter at 1.7 m above ground
were measured with a diameter tape and mapped in a
1.44-ha plot (Fujimori 1977). Allometric equations for
tree height and stem volume were developed for the
site. Equations for tree height as a function of diam-
eter were developed from a subsample of live trees
measured using an Abney level. An equation for
Sequoia stem volume as a function of diameter and
height was developed from a sample of 8 recently
fallen trees (see Appendix). In 2001, we measured
diameter on all live trees within the plot and noted tree
ingrowth and mortality by species since 1972.
Using the 1972 stem map, diameter growth of
individual live trees over the 29-yr period was
determined. Mean annual diameter increments of

each species were calculated. To account for effects
of tree size and canopy position on growth, Sequoia
diameter increments were calculated for each of the
three canopy strata recognized by Fujimori (1977).
These included a lower stratum (diameter <30 cm), a
middle stratum (diameter 30–114 cm), and an upper
stratum (diameter >114 cm). Sequoia tree heights in
2001 were estimated from diameter using one of the
stratum-specific allometric equations (Fujimori
1977).
Estimation of tree ANPP followed the general for-
mula summarized by Newbould (1967), Ogawa
(1977) and Long (1982) as the total annual growth of:
1) live trees, 2) trees or tree parts lost to mortality, and
3) tree parts consumed. Tree parts shed or consumed
were ignored, however. Stand volume increment since
1972 was initially calculated with ingrowth and
mortality components. Whenever possible, the mea-
sured diameter increment for each tree was used. If a
measured Sequoia increment was uncertain (because
of measurement location or tree identity), the mean
increment for the corresponding canopy stratum (see
above) was applied. Trees that died over the 29-yr
interval were assigned a diameter increment equal to
half the corresponding mean increment. Diameter
increments of ingrowth trees were calculated using an
initial diameter of 10 cm. Height increments were
calculated from diameter measurements using a stra-
tum-specific allometric equation (Fujimori 1977). For
trees with a measured height value in 1972, the
increment was added to the measured height. Para-
bolic volume (see Appendix) was calculated for 1)
live trees in 1972, 2) live trees in 2001, and 3) live
trees in 2001 plus trees that died over the interval.
Annual parabolic volume increment was then calcu-
lated with and without compensation for mortality.
Stem volume of Sequoia was estimated using three
methods (see Appendix): 1) the allometric relation
developed by Fujimori (1977) for the study site, 2) a
second allometric relation developed using stem taper
data on hundreds of old-growth trees in the region
(Parks 1952), and 3) the estimation ratios of Westman
and Whittaker (1975). The latter method involved
multiplying the parabolic volume by 0.9. Stem mass
estimates were obtained by multiplying the stem
volume by the specific gravity. Branch biomass of
Sequoia was estimated as 0.12 of the stem mass
(Westman and Whittaker 1975).
Stem NPP of Sequoia was calculated as the annual
stem volume increment times the specific gravity. A
range of branch NPP values was obtained by multi-
179
plying the stem NPP by 0.25 and by 0.30 (Westman
and Whittaker 1975). A range of twig and leaf NPP
values was estimated as the combined stem and
branch NPP multiplied by 0.3 and by 0.5.
Separate biomass and production estimates were
made using two different specific gravity values for
Sequoia. Westman and Whittaker’s (1975) estimates
were based on an intermediate value of 0.33. How-
ever, Green et al. (1999) note that 0.38 is appropriate
for old-growth Sequoia. Some of our dead wood
samples (see below) also indicated that 0.38 best
represented the study forest. So, for comparative
purposes we used 0.33, but we also used 0.38 and we
considered this to be a more accurate value for the
study forest. Other species comprised a very small
proportion of the stand biomass and NPP. Total
aboveground values for these species were estimated
directly from parabolic volume and specific gravities
of each species.
Woody detritus
Woody detritus volume in 2001 was estimated with
plot measurements for standing dead trees (or snags)
and line transects for logs. Volume of standing dead
trees (>1.7 m tall) within the 1.44-ha plot was cal-
culated from measurements of diameter and height.
The volumetric shape (cylindric, parabolic, or conic)
and stage of wood decay (stage 1–5) of each standing
dead tree were noted. Down dead wood was sampled
with eight 100-m transects across the study forest. To
avoid transect orientation bias, four of the transects
ran roughly north to south and the other four ran east
to west. The lateral distance between parallel transects
exceeded 100 m. Fine dead wood (<10 cm diameter at
intercept) was tallied along transect segments in size
classes. The three smallest size classes followed those
of Brown (1974). Class 1 items were <0.7 cm dia-
meter, class 2 items were 0.7 to 2.5 cm diameter, and
class 3 items were 2.6 to 7.6 cm diameter. A fourth
fine wood class included items 7.7 to 9.9 cm diameter.
Items in classes 1 and 2 were tallied on one 7-m
segment per 100-m transect. Items in classes 3 and 4
were tallied on one 14-m segment per 100-m transect.
Coarse dead wood ( £ 10 cm diameter at intercept)
was measured for diameter at point of intercept along
each 100-m transect, identified to species, and
assessed for stage of decay (stage 1
5; Sollins 1982).
Specific gravity of Sequoia wood by stage of decay
was estimated with wood samples measured for
180
volume and dry weight. For each stage of decay a set
of samples, including representative proportions of
sapwood and heartwood, was collected from the study
site. Most items were cylindric and volume could be
estimated from length and diameter. Other items were
submersed in water and the displaced volume was
measured. All samples were then oven dried at 55 C
(to allow nutrient analysis) until constant dry weight
was obtained (ca. 2 weeks). A subset of these samples
was subsequently dried at 105 C for two additional
days and the dry weight of all samples was adjusted
using the mean percentage of weight lost at the higher
temperature.
Rate of coarse woody detritus input was determined
from tree mortality in the 1.44-ha plot over a 29-yr
period. Stem volume of the dead individuals was
estimated using the two allometric equations
described above. The parabolic approach of Westman
and Whittaker (1975) was not used in the estimation
of dead tree volumes.
Volume of fallen woody detritus was calculated
using Van Wagner’s (1968) formula (see Appendix).
Mass of coarse detritus was estimated for each stage
of decay using our specific gravities. Fine woody
detritus calculations followed those of Brown (1974);
a representative diameter was used for each size class
and a correction factor for intercept bias of nonhori-
zontal pieces was applied to the three smallest diam-
eter classes (see Appendix). A specific gravity of 0.33
was used for all fine detritus mass estimates.
Some authors include dead material as a biomass
component (e.g., Sprugel 1985). We retain the tradi-
tional definition of biomass as the weight of material
in living trees (Ricklefs 1973) and refer to material
from dead trees and fallen parts as detritus.
Results
Biomass
Total Sequoia stem biomass estimates for the study
plot in 1972 ranged from 2989 to 5133 Mg ha
1.
Estimates depended on the method of stem volume
estimation and the specific gravity value applied
(Table 1, Table 2). Both allometric relations gave
lower stem volume estimates than the parabolic esti-
mates of Westman and Whittaker (1975). Comparison
of estimated stem volumes to independent data on
measured volumes of large tree stems (Sawyer et al.
2000a) supported the validity of the allometric rela-
tions (Figure 1). Both allometric relations generated
mean values within one standard deviation (of the
estimate) of the mean measured value. By contrast,
the parabolic estimates were much greater, overesti-
mating the measured value in all cases. Fujimori’s
allometric relation overestimated the measured stem
volume in 83% of the cases, while the second allo-
metric relation underestimated the measured valued in
67% of the cases. We considered our stem biomass
estimates based on the allometric relations (and a
specific gravity of 0.38) to be more accurate than any
of the parabolic estimates. These relatively accurate
estimates ranged from 3442 to 4143 Mg ha
1.
Sequoia branch biomass estimates in 1972 ranged
from 359 to 616 Mg ha
1 (Table 1, Table 2). Esti-
mates of 413 to 497 Mg ha
1 corresponded to our
stem biomass estimates of 3442 to 4143 Mg ha
1
above. Combining stem and branch compartments we
obtained total Sequoia aboveground biomass esti-
mates of 3855 to 4640 Mg ha
1. We considered these
estimates to be relatively accurate. It should be noted,
however, that our total Sequoia biomass estimates,
calculated using a variety of methods, had a broader
range (3348 to 5749 Mg ha
1; Table 1, Table 2). Total
stand biomass was only slightly greater, ranging from
3350 to 5751 Mg ha
1.
Sequoia stem biomass declined 7% over the 29-yr
period. Although biomass declined over the 1.44-ha
area taken as a whole, some patches within the study
plot increased in biomass. An examination of biomass
change against grain size (the smallest area of obser-
vation and measurement; sensu Wiens 1989) revealed
that patches exhibiting biomass increases were in the
0.01 to 0.4 ha size range. All larger patches or col-
lections of patches decreased in biomass. Plotting
temporal variance in stem biomass against grain size
(e.g., Levin 1992) revealed high variance at grain
sizes less than 0.05 ha (Figure 2a). The log-log rela-
tionship was approximately linear (Figure 2b).
Production
Sequoia stem ANPP for our study site ranged from 3 to
7 Mg ha
1yr
1 (Table 1, Table 2). Estimates based on
the allometric relations for stem volume were lower
than those based on parabolic volume and estimation
ratios (Westman and Whittaker 1975). Consideration
of trees lost to mortality resulted in slightly greater
ANPP values (Table 1, Table 2). The use of a relatively
large specific gravity of 0.38 for Sequoia also elevated
 181

Table 1. Comparison of biomass and production estimates for Sequoia sempervirens forests using a specific gravity of 0.33 for Sequoia.
Volume, biomass, basal area, density and height values are reported for the initial sampling date (1972 for the current study). Values in
parentheses include contributions of trees dying over the course of the 29-yr study. Means or ranges of estimates from previous studies are based
on Whittaker (1966), and Westman and Whittaker (1975). Allometric estimates 1 and 2 refer to stem volume equations based on Fujimori (1977)
and Parks (1952), respectively.

Parameter This study Previous studies

Terrain Alluvial flat Alluvial flat Slope

Elevation (m) 80 60-240 50-270
Distance from coast (km) 25 20-30 2-20
Slope and aspect level level mixed
Total sample area (ha) 1.44 0.8 0.3
Density (stems ha
1)
Sequoia 310 280 330
Other species 70 60 590
Weighted mean height (m) 91 80 46
Mean radial increment (mm yr
1)
All trees 0.75 0.86 0.95
All Sequoia trees 0.80 – –
Lower stratum 0.67 – –
Middle stratum 0.54 – –
Upper stratum 0.97 – –
Basal area (m2 ha
1)
Sequoia 329 248 112
Other species <1 <1 22
All species 330 248 134
Basal area increment (m2 ha
1 yr
1)
Sequoia 0.388 (0.404) – –
Other species 0.004 (0.005) – –
Parabolic volume increment (cm3 m
2 yr
1)
Sequoia 1816 (1883) 1586 850
Other species 3 (4) 1224 232
Sequoia stem vol. increment (cm3 m
2yr
1)
Allometric estimate 1 1299 (1348) – –
Allometric estimate 2 1047 (1078) – –
Parabolic volume (m3 ha
1)
Sequoia 15009 9195 2818
Other species 4 <1 288
Sequoia stem volume (m3 ha
1)
A1) Parabolic volume X 0.9 13508 – –
B1) Allometric estimate 1 10903 – –
C1) Allometric estimate 2 9059 – –
Sequoia stem biomass (Mg ha
1)
A2) A1 X 0.33 4458 – –
B2) B1 X 0.33 3598 – –
C2) C1 X 0.33 2989 – –
Sequoia branch biomass (Mg ha
1)
A3) A2 X 0.12 535 – –
B3) B2 X 0.12 432 – –
C3) C2 X 0.12 359 – –
Sequoia stem ANPP (g m
2 yr
1)
A4) Parabolic stem vol. incr. X 0.33 599 (621) – –
B4) (Allometric stem vol. incr. 1) X 0.33 429 (445) – –
C4) (Allometric stem vol. incr. 2) X 0.33 346 (356) – –
Sequoia branch ANPP (g m
2yr
1)
A5) A4 X 0.25 to 0.3 150-180 (155-186) – –
B5) B4 X 0.25 to 0.3 107-129 (111-134) – –
C5) C4 X 0.25 to 0.3 87-104 (89-107) – –
Sequoia stem and branch ANPP (g m
2 yr
1)
A6) A4+A5 749-779 (776-807) – –
182

Table 1. Continued.

Parameter This study Previous studies

B6) B4+B5 536-558 (556-579) – –
C6) C4+C5 433-450 (445-463) – –
Sequoia twig and leaf ANPP (g m
2 yr
1)
A7) A6 X 0.3 to 0.5 225-390 (233-404) – –
B7) B6 X 0.3 to 0.5 161-279 (167-290) – –
C7) C6 X 0.3 to 0.5 130-225 (134-232) – –
Total ANPP (g m
2 yr
1)
Sequoia
A8) A6+A7 974-1169 (1009-1211) – –
B8) B6+B7 697-837 (723-869) – –
C8) C6+C7 563-675 (579-695) – –
Other tree species 1 (2) – –
All tree species
A9) Parabolic estimate 975-1170 (1011-1213) 1255 1238
B9) Allometric estimate 1 698-838 (725-871) – –
C9) Allometric estimate 2 564-676 (581-697) – –
Total tree biomass (Mg ha
1)
Parabolic estimate 4995 1155 3069
Allometric estimate 1 4032 – –
Allometric estimate 2 3350 – –

ANPP estimates. We consider our stem ANPP
estimates of 4 to 5 Mg ha
1yr
1 to be the most accurate
(Table 2). They are based on the allometric relations
and a specific gravity of 0.38. Also, they include con-
tributions of trees dying over the 29-yr study period.
Ranges of Sequoia branch ANPP, estimated using the
ratios of Westman and Whittaker (1975), were 1 to 2 Mg
ha
1yr
1 (Table 1, Table 2). Those for twig and leaf
components were 1 to 5 Mg ha
1yr
1 (Table 1, Table 2).
Total tree ANPP estimates ranged from 6 to 14 Mg
ha
1yr
1 in our study (Table 1, Table 2). We consid-
ered our estimates of 7 to 10 Mg ha
1yr
1 to be
comparatively accurate. They were based on the allo-
metric relations and a specific gravity of 0.38 (Table 2).
depending on the allometric relation used to estimate
stem volume (see methods). Annual input averaged
5.7 to 6.9 Mg ha
1 (Table 3). Assuming constant
input and steady state total mass, a decay rate constant
of 0.22 to 0.026 was obtained by dividing the annual
input by the total mass of coarse detritus (Jenny et al.
1949; Olson 1963; Whittaker 1975; Sollins 1982;
Onega and Eickmeier 1991). The total volume of fine
woody detritus on the forest floor was 16 m3 ha
1
(Table 3). Total mass in this compartment was esti-
mated at 5 Mg ha
1.
Discussion

Woody detritus Biomass and patch dynamics

The total volume of coarse woody detritus was 797
m3 ha
1 (Table 3). The total mass was 262 Mg ha
1.
Fallen wood comprised most of the coarse woody
detritus. The log component was 743 m3 ha
1 (242
Mg ha
1). The snag component was 54 m3 ha
1 (20
Mg ha
1). Snag basal area and density were 12.7 m2
ha
1 and 12 stems ha
1, respectively. For comparison,
live tree basal area and density in 2001 were 325 m2
ha
1 and 170 stems ha
1, respectively.
The input of coarse woody detritus in the study plot
over 29 yr was 437 to 522 m3ha
1 (166-199 Mg ha
1),
The slight decline (7%) in biomass during the three-
decade study supports other evidence that the stand is
in an advanced stage of development. Rapid, stand-
wide biomass accumulation typical of early devel-
opment (Bormann and Likens 1979; Peet 1981) has
clearly ceased. The stand has reached the old growth
stage or is in transition toward old growth (Oliver
1981; Peet and Christensen 1987). Large tree boles
(both live and dead), characteristic of old-growth
coniferous forest structure (Franklin et al. 1981), are
abundant. Trees greater than 1000 years of age are
 183

Table 2. Biomass and production estimates for the study site using a specific gravity of 0.38 for Sequoia. Volume and biomass values are
reported for the initial sampling date (1972). Allometric estimates 1 and 2 refer to stem volume equations based on Fujimori (1977) and Parks
(1952), respectively. Values in parentheses include contributions of trees dying over the course of the 29-yr study. See Table 1 for stem volume
increments.

Parameter Value

Sequoia stem volume (m3 ha
1)

A1) Parabolic volume X 0.9 13508
B1) Allometric estimate 1 10903
C1) Allometric estimate 2 9059
Sequoia stem biomass (Mg ha
1)
A2) A1 X 0.38 5133
B2) B1 X 0.38 4143
C2) C1 X 0.38 3442
Sequoia branch biomass (Mg ha
1)
A3) A2 X 0.12 616
B3) B2 X 0.12 497
C3) C2 X 0.12 413
Sequoia stem ANPP (g m
2 yr
1)
A4) Parabolic stem vol. incr. X 0.38 690 (716)
B4) (Allometric stem vol. incr. 1) X 0.38 494 (512)
C4) (Allometric stem vol. incr. 2) X 0.38 399 (410)
Sequoia branch ANPP (g m
2 yr
1)
A5) A4 X 0.25 to 0.3 173-207 (179-215)
B5) B4 X 0.25 to 0.3 124-148 (128-154)
C5) C4 X 0.25 to 0.3 100-120 (103-123)
Sequoia stem and branch ANPP (g m
2 yr
1)
A6) A4+A5 863-897 (895-931)
B6) B4+B5 618-642 (640-666)
C6) C4+C5 499-519 (513-533)
Sequoia twig and leaf ANPP (g m
2 yr
1)
A7) A6 X 0.3 to 0.5 259-449 (269-466)
B7) B6 X 0.3 to 0.5 185-321 (192-333)
C7) C6 X 0.3 to 0.5 150-260 (154-267)
Total ANPP (g m
2 yr
1)
Sequoia
A8) A6+A7 1122-1346 (1164-1397)
B8) B6+B7 803-963 (832-999)
C8) C6+C7 649-779 (667-800)
Other tree species 1 (2)
All tree species
A9) Parabolic estimate 1123-1347 (1166-1399)
B9) Allometric estimate 1 804-964 (834-1001)
C9) Allometric estimate 2 650-780 (669-802)
Total tree biomass (Mg ha
1)
Parabolic estimate 5751
Allometric estimate 1 4642
Allometric estimate 2 3857

not uncommon (Fujimori 1977), indicating that at
least a millennium has passed since the last stand-
replacing disturbance.
Although pervasive disturbances such as fire and
flooding affect the dynamics of riparian Sequoia for-
ests (Stone and Vasey 1968;Veirs 1982; Agee 1993;
Sawyer et al. 2000a), the predominant form of dis-
turbance in our study forest over the last three decades
has been tree-fall gaps created by trees without severe
fire injury. Our analysis of temporal variance in bio-
mass by grain size, showing greater variance at grain
sizes less than 0.05 ha, shows the effect of scattered
mortality of individual large trees. Recent canopy
gaps in our study plot are generally less than 0.05 ha;
however, a survey of the study area revealed gaps
approaching 0.2 ha in size (Busing and Fujimori
184

Figure 1. Comparison of calculated Sequoia stem volumes against measured volumes. Means and standard deviations for each method are
shown. Volume data in the ‘measured’ category are from Sawyer et al. (2000a). They consist of very large trees (87–110 m tall; n ¼ 12) within
the region that were measured in the field. All other categories displayed are calculated volume estimates applying different formulas based on
tree diameter and height. The various formulas range from simple parabolic volume estimates (Westman and Whittaker 1975) to allometric
volume estimates developed with field data and regression analysis (Fujimori 1977).

2002). Further analysis of biomass variance by grain
size reveals a log-log linear relationship. Levin (1989)
states that departure from a linear relationship indi-
cates ‘...operation of distinct mechanisms on different
scales.’ For biomass dynamics, it appears that distinct
mechanisms of this sort have not operated during the
course of this study.
A general conclusion from the analysis of biomass
dynamics is that small-patch dynamics are significant.
A shifting-mosaic of small patches influenced by tree
falls is characteristic of many old-growth forests
(Bormann and Likens 1979; Pickett and White 1985;
Busing 1998; Fujimori 2001), including those domi-
nated by conifers (McCarthy 2001). Our short glimpse
at biomass dynamics within a stand suggests that this
type of small patch dynamics can play a substantial
role in old-growth Sequoia forests as well.
Production
Our ANPP estimates of 6 to 14 Mg ha
1yr
1 fall
within the range of values reported for other temper-
ate forests. Our estimates tend to be less than the
Sequoia forest ANPP means from Westman and
Whittaker (1975). One possible explanation is simply
that our study stand has slower growth. Indeed, our
mean radial increments were slightly lower. However,
the primary reason for the discrepancy is that the al-
lometric equations we use to estimate stem volume
give lower values. Even our estimates using a larger
Sequoia specific gravity and including ANPP of trees
lost to mortality tend to be lower than those of
Westman and Whittaker (1975) when allometric stem
volumes are used.
We consider our ANPP estimates of 7 to 10 Mg
ha
1yr
1 to be the most accurate. They are based on
allometric stem volume and a specific gravity of 0.38.
They also include contributions of trees dying over
the course of the study. These ANPP values are
slightly below average for old temperate forests
(Whittaker 1966; Fujimori 2001), but similar to those
reported for other old coniferous forests of the Pacific
Slope of North America (Long 1982). Our estimates
do support the idea that old Sequoia forests are not
highly productive (Whittaker 1966; Westman and
Whittaker 1975). Even though biomass exceeds that
of most other temperate forests by a factor of ten or
more, our ANPP estimates are not high.
The ratio of biomass to annual NPP is very large
(ca. 400) for our study forest. Whittaker (1975)
termed this the biomass accumulation ratio. It typi-
cally ranges from 20 to 50 in mature forests. The
extreme biomass accumulation ratio value from our
study is a result of the exceptionally large biomass, as
ANPP is moderate.
The fact that our study stand is old and strongly
dominated by Sequoia deserves consideration in
 185

Table 3. Volume and mass of woody detritus at the study site. The
minimum diameter of coarse woody detritus is 10 cm. Decay classes
range from undecayed (class 1) to well decayed (class 5).

Parameter Volume (m3ha–1) Mass (Mg ha–1)

Coarse woody detritus

Fallen, decay class 1 121 46
Fallen, decay class 2 98 36
Fallen, decay class 3 451 149
Fallen, decay class 4 67 11
Fallen, decay class 5 6 1
Fallen total 743 242
Standing 54 20
Annual input 15–18 5.7–6.9
Fine woody detritus
Fallen (<0.7 cm diam.) 0.4 0.1
Fallen (0.7-2.5 cm diam.) 2.3 1
Fallen (2.6-7.6 cm diam.) 6.9 2
Fallen (7.7-9.9 cm diam.) 6.3 2
Fallen total 16 5

Mg ha
1yr
1 have even lower mean ANPP per stem

(<0.014 Mg stem
1yr
1; Busing et al. 1993; Busing
1998). Radial growth of old Sequoia trees is relatively
slow, but individual overstory trees do tend to have
greater ANPP than trees of other species in the region
and in other temperate forests. Nonetheless, ANPP in
old, pure stands of Sequoia is not above average for
temperate forests.

Woody detritus

The volume and mass of coarse woody detritus (797

Figure 2. Temporal variance in biomass versus grain size (or area of
observation) over the three-decade period. Variance is plotted against
grain size using: (a) arithmetic axes, and (b) logarithmic axes.
interpretation of our ANPP results. Clearly, younger
stands of Sequoia can have much greater ANPP (e.g.,
Hallin 1934; Fritz 1945; Westman and Whittaker
1975; Olson et al. 1990). Also, data presented by
Westman and Whittaker (1975) suggest that mixed-
species stands containing Sequoia tend to be more
productive. Even though mean ANPP per stem (>2
cm diameter) is greater in pure, old Sequoia stands
(>0.02 Mg stem
1yr
1) than in mixed-species stands
with Sequoia (<0.015 Mg stem
1yr
1; Westman and
Whittaker 1975), old, pure stands have lower stem
densities and ANPP. By comparison, old temperate
deciduous stands having ANPP ranging from 6 to 12
m3 ha
1 and 262 Mg ha
1) are much greater than
average for temperate forests. Yet, our values are
within the range of those reported for Sequoia forests
(Sawyer et al. 2000b) and for other old coniferous
forests of the Pacific Slope of North America
(Harmon et al. 1986). It has been speculated that
coarse woody detritus of Sequoia stands may be
consumed by fires (Agee 1993) or buried by flood
sediments on certain sites (Sawyer et al. 2000a). In
our study forest, however, there was no evidence of
recent burning or burial of coarse woody detritus.
Heavily decayed logs of Sequoia were common. No
charred fallen detritus was observed. Likewise, sub-
stantial burial of coarse detritus was not evident. We
conclude that input and decay processes regulate the
coarse woody detritus pool in our study forest, al-
though this was probably untrue a century ago before
fire suppression. Coarse detritus input, estimated from
29 years of tree mortality over 1.44 ha, is 5.7 to 6.9
186
Mg ha
1yr
1. These values are well above average for
temperate forests, but within the range reported for old
coniferous forests of the Pacific Slope (Harmon et al.
1986). Dividing our estimates of annual coarse detri-
tus input by the total amount of coarse detritus, we
obtain decay rate constants from 0.022 to 0.026.
These estimates fall within the range of other values
for temperate coniferous forests (Stokland 2001).
Assuming negative exponential decay (Olson 1963),
they project 50% decay in 27 to 32 years and 95%
decay in 115 to 136 years. Thus, we suggest that the
decay rate at our study site is not extremely slow. It is
below average for forest ecosystems in general, but it
appears to resemble that of other temperate coniferous
forests.
Relationships of coarse woody detritus to biomass
and ANPP can be characterized for purposes of
comparison and estimation. The ratio of coarse
detritus mass to biomass is 0.06 to 0.07. These values
are within the range reported for temperate and trop-
ical forests (Muller and Liu 1991; Edwards and Grubb
1977). They tend to be lower than values calculated
for other coniferous forests of the Pacific Slope,
however. The ratio of annual coarse detritus input to
biomass ranges from 0.001 to 0.002. These compar-
atively low values reflect the low mortality rate of
Sequoia (Busing and Fujimori 2002). The annual
coarse detritus input rate divided by annual ANPP is
approximately 0.8. The comparable ratio for stem
ANPP is approximately 1.4. Given that a vast pro-
portion of coarse detritus is stem wood, these results
are in agreement with the decline in live biomass
observed over the course of this study.
Conclusions
The biomass of our study stand is more than ten times
the average for temperate coniferous forests (Whit-
taker and Likens 1975). Yet, ANPP is moderate and
woody detritus levels are not extreme. Thus, biomass
accretion and loss are not rapid in this old-growth
forest. Population processes of Sequoia, the dominant
species, partly explain these observations. Individual
tree boles in the stand approach 4 m in diameter and
100 m in height (Fujimori 1977), but diameter growth
tends to be slow. In addition, the Sequoia population
is characterized by very low rates of tree ingrowth and
mortality at the old-growth stage (Busing and Fuji-
mori 2002). Low to moderate rates of biomass
accretion and loss reflect the slow canopy turnover of
this Sequoia dominated forest.
Acknowledgements
We are grateful to Bruce Caldwell and Tom Hayes for
use of laboratory space and equipment for analysis of
our wood samples. Professor R.H. Waring provided a
helpful discussion on estimating production in wes-
tern conifers. Steve Hostetler gave the manuscript a
careful proofreading. Several anonymous reviewers
made constructive comments on the manuscript. This
study was initiated while the first author was at the
Corvallis Forestry Sciences Lab and Oregon State
University.
Appendix
The mathematical formulas used in volume calcula-
tions for trees and woody detritus are described below.
Pertinent information regarding the formulas and their
application is provided.
Tree stem volume was estimated from tree diameter
and height using three separate equations. The first
equation relies on the assumption that stem volume is
a simple function of the parabolic volume:
V = p · (D/2)2 · H · 0.5
Where V is volume (m3), D is bole diameter (m),
and H is tree height (m). Following Westman and
Whittaker (1975) we multiplied the parabolic volume
by 0.9 to obtain an estimate of Sequoia stem volume.
The second equation for stem volume (allometric
volume estimate 1 herein) was developed using a
sample of 8 Sequoia trees from the study stand. Fu-
jimori (1977) obtained the following relation from
linear regression of logarithmically transformed vari-
ables:
Log10 V = 0.9784 · (Log10 D2H)
0.4843
The regression R2 was 0.99 and the standard error of
the estimate was 0.029. Standard errors for the inter-
cept and the slope were 0.12 and 0.043, respectively.
The third equation for stem volume (allometric
volume estimate 2 herein) was developed using a set
of 9 Sequoia trees with typical bole taper. Taper
profiles were obtained from summaries of hundreds of
old-growth forest trees in the study region (Parks
1952). We obtained the following relation from linear
regression of logarithmically transformed variables:

2 structure in an old Sequoia sempervirens forest. Journal of Veg-
Log10 V = 0.9246 · (Log10 D H)
0.4147
The regression R2 was 0.99 and the standard error of
the estimate was 0.028. Standard errors for the inter-
cept and the slope were 0.046 and 0.020, respectively.
Volume of fallen woody detritus was estimated
using line intercept methods. Coarse woody detritus
(‡10 cm diameter) on or near the forest floor was
estimated from piece diameter at point of intercept
using the formula of VanWagner (1968):
V = p2 · R(D2/8L)
Where V is volume (m3m
2), D is piece diameter
(m), and L is total transect length (m).
Volume of fine woody detritus (<10 cm diameter)
on the forest floor was estimated by size-class using a
modification of VanWagner’s formula:
V = p2 · N · A ·(Q2/8L)
Where N is the number of pieces tallied, A is the
correction factor for non-horizontal pieces, and Q is
the quadratic mean diameter (m) of pieces in the size
class (Brown 1974). Values of A were set to 1.13 for
all size classes having pieces £ 7.6 cm diameter
(Brown 1974). Otherwise the value of A was 1.0.
Values of Q followed Brown’s (1974) composite
values for conifers: 0.00312 for pieces <0.7 cm dia-
meter, 0.0137 for pieces 0.7 to 2.5 cm diameter,
0.0422 for pieces 2.6 to 7.6 cm diameter, and 0.0843
for pieces 7.7 to 9.9 cm diameter.
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