1429
Growth and foliar nutrient concentrations of Pacific silver fir
M. A. RADWAN, M. D. MURRAY, AND J. M. KRAFT
USDA Forest Service, Forestry Sciences Laboratory, Pacific Northwest Research Station,
3625 93rd Avenue, SW, Olympia, WA 98502, U.S.A.
Received March 22, 1989
Accepted July 4, 1989
RADWAN,
M. A., MURRAY, M. D., and KRAFT, J. M. 1989. Growth and foliar nutrient concentrations of Pacific silver
fir. Can. J. For. Res. 19: 1429-1435.
Nineteen natural stands of Pacific silver fir (Abies amabilis (Doug!.) Forbes) were assessed for selected site and tree
characteristics. The stands, located throughout western Washington, were 26 to 37 years old;<they occurred at various
elevations (110-1300 m) and on different soil types, and varied greatly in associated plant species. Site index at 50 years
breast-high age ranged from 12 to 24 m. Terminal growth per year during the previous 3 years, needle number per
centimetre of twig tissue, needle weight and area, and foliar concentrations of 13 elements varied among the 19 stands.
Compared with associated western conifers, silver fir had moderate terminal growth, larger and heavier needles, more
needles per centimetre of twig tissue, and lower foliar concentrations of N, S, and Mg. There were many significant
correlations among the different variables. The foliar nutrient data obtained provide base-line information for future
research in nutrition and fertilization of silver fir. These data and the significant correlations between nutrients and
site-stand characteristics suggest that U) silver fir may be more efficient than some western conifers in use of some
nutrients, (ii) productivity of silver fir may be improved by application of certain fertilizers; fertilization may be
particularly effective dUring the seedling stage when growth is typically slow, and (iii) levels of some foliar nutrients
may be useful as indicators of site quality for silver fir production.
RADWAN, M. A., MURRAY, M. D., et KRAFT, J. M. 1989. Growth and foliar nutrient concentrations of Pacific silver
fir. Can. J. For. Res. 19: 1429-1435.
Dix-neuf peuplements naturels de Sapin gracieux (Abies amabilis (Doug!.) Forbes) furent seiectionnes pour Ie type
de station et les caracteristiques des arbres. Les peuplements,
de 26 a 37 ans, etaient disperses dans I'ouest de
I'etat de Washington. Ils etaient situes it diverses altitudes (110 it 1300 m) et sur differents types de sol, et comportaient
une grande variete d'especes associees. L'indice de station it 50 ans (age it hauteur de poitrine) allait de 12 it 24 m.
La croissance apicale des 3 annees precedentes, Ie nombre d'aiguilles par centimetre de rameau, Ie poids et la surface
des aiguilles, et la concentration foliaire de 13 elements variaient d'un peuplement it I'autre. Compare aux coniferes
de I'ouest auxquels il est associe, Ie Sapin gracieux avait une croissance apicale moyenne, des aiguilles plus grosses
et plus lourdes, plus d'aiguilles par centimetre de rameau, et une plus faible concentration de N, S et Mg. Plusieurs
variables etaient significativement correlees entre elles. Les donnees recueillies sur la concentration foliaire des elements
nutritifs constituent de I'information de base pour de futurs travaux de recherche sur la nutrition et la fertilisation
du Sapin gracieux. Ces donnees et les correlations significatives entre les elements nutritifs et les caracteristiques des
stations et des peuplements suggerent (I) que Ie Sapin gracieux peut etre plus efficace que certains coniferes de l'ouest
dans I'utilisation de certains elemens nutritifs, (ii) qu'il est possible d'ameliorer la productivite du Sapin gracieux par
I'application de fertilisants qui pourraient etre particulierement efficaces au stade de semis lorsque la croissance est
typiquement faible, et (iii) que la concentration de certains elements nutritifs pourrait servir d'indice de qualite de
station pour la production du Sapin gracieux.
[Traduit par la revue]
Introduction
Pacific silver fir (Abies amabilis (Doug!.) Forbes) is an
important softwood tree species in the forests of north­
western North America. In the Pacific Northwest, silver fir
is the most important of the western true firs (Abies spp.)
(Franklin 1982). Silver fir has a natural range exteilding from
the southeastern edge of Alaska to northwestern California,
at elevations ranging from sea level to over 1600 m (Fowells
1965). In Canada, silver fir is found mainly in the Coast
Mountains and islands of British Columbia (Fowells 1965).
In Washington State, United States, silver fir occurs
throughout the Cascade Range, the Olympic Mountains, and
the Coast Ranges of southwestern Washington (Fowells
1965; Murray and Treat 1980; Packee et al. 1982). In these
and other areas throughout its range, silver fir occurs most
frequently in mixture with other tree species. The most com­
mon associated conifer in. western Washington is western
hemlock (Tsuga heterophylla (Raf.) Sarg.).
Currently there is much interest in the intensive manage­
ment of silver fir, especially at elevations of 700 to 1400 m.
Like other true firs in the West, silver fir has become imporPrinlcd in Canada I Imprimi: au Canada
tant as a forest resource as forest managers move to higher
elevations in search for new wood supplies. At such eleva­
tions, Dimock (1958) suggested that in some overmature
climax types, net yields of unmanaged silver fir stands may
equal those of western hemlock. Others (Packee et al. 1982)
3
reported yields of 950 to 1600 m /ha for silver fir
dominated stands at 100 to 125 years of age.
Prescribing effective management practices to increase
productivity of silver fir requires much basic information
on the biology and nutrition of the species. To date,
however, relevant knowledge lags far behind management
needs. For example, review of the literature shows that there
are no guidelines for fertilization of silver fir; only one study
dealing specifically with effect of fertilization on growth of
the species (Gallager 1964) has been conducted. Similarly,
reports on the foliar nutrient content of the trees in natural
stands (Beaton et al. 1965b; Gessel and Orians 1967; Turner
and Singer 1976; Will and Youngberg 1979; Gessel and
Klock 1982; Cochran et al. 1986) were very limited in scope.
Each of these reports involved only one site, and none
provided information on concentrations of all the essential
CAN. J. FOR. RES. VOL. 19, 1989
1430
nutrient elements or on the deficiency and toxicity levels of
any nutrient. Furthermore, there are no data in the literature
on the relationships among the foliar elements or between
the elements and site and stand characteristics.
This study was conducted to (i) determine site quality,
I
51
100 km
4
• Bellingham
2,.3
1
growth, and foliar chemical composition of silver fir from
different stands in western Washington, (ii) assess the
relationships between the different variables determined, and
(iii) provide base-line information for future nutritional
work.
Materials and methods
Stand and tree selection
Nineteen natural stands of Pacific silver fir were selected for
investigation. The stands are located in western Washington, at
elevations ranging from 110 to 1300 m (Fig. 1). Stands were selected
to assure representation of ages economically suitable for fertiliza­
.Olympia
13
12
·9
•
10 •·
tion and a wide range of soil and site conditions. Each stand had
an area of at least 0.5 ha where topography and stand structure
were fairly uniform and silver fir was at least 20010 of the basal area.
In each stand, eight dominant or codominant trees were ran­
domly selected for study. Selected trees ranged in age from 26 to
37 years. All trees appeared healthy with no evidence of previous
top damage.
Estimation oj site index
At each location, selected trees were felled during October and
November 1985. Tree heights and ages were determined, and site
index at 50 years (total age) was estimated according to Hegyi et aL
(1981).
Height-growth measurements
Terminal growth of the test trees for the 3 years immediately
before felling was measured, and average height growth per year
was computed.
Sampling joliage
Immediately after feIling, one composite, representative sample
of about 200 g (fresh weight) of foliage was collected from the eight
trees of each stand. Sampling was limited to vigorous branches
in the sunlit portion of the crown. Each sample consisted of 5-cm
tips of the current year's growth of secondary laterals cut from
all sides of the trees. Samples were individually placed in precooled
glass containers and transported to the laboratory in a portable
cooler.
Determination oj needle numbers, areas, and weights
FIG. 1. Approximate geographic location of study sites in
western Washington, United States. Site descriptions are in Table 1.
Total N of the ovendried needles was determined by the standard
micro-Kjeldahl procedure (Bremner and Mulvaney 1982). Analyses
of other elements were performed on the dry needles or on solu­
tions of their ash as follows: P by the molybdenum blue technique
and Mo by the thiocyanate colorimetric method (Chapman and
Pratt 1961); total S by a turbidimetric technique (Butters and
Chenery 1959); B by the quinalizarin procedure (Horwitz 1980);
and Ca, Mg, K, Fe, Mn, Cu, Zn, and AI by standard atomic
absorption spectrophotometric technique (Perkin-Elmer Corpora­
tion 1976). All analyses were carried out at our laboratory.
Statistical analysis
Correlation coefficients (r) between the variables measured were
calculated using each site as an individual observation (Snedecor
1961). Differences between individual sites could not be statistically
tested because observations were made on subsamplesj no "true"
replication was possible at the different sites.
Two subsamples of three lateral twigs each were taken at random
from each composite sample. Needles were separated from stems
and counted. Stem length was measured, and average number of
needles per centimetre of twig length was calculated. Area of the
separated needles was determined using a LI-COR, LI-3100 area
meter. I Needles on remaining twigs of each composite sample
were also separated from stems and buds, and needles of each
sample were thoroughly mixed. Two subsamples of 200 needles
each were taken at random from each composite needle sample.
Needles were dried to constant weight at 65°C, and average weights
of individual needles were calculated for each site.
Chemical analysis
Separated needles, remaining after samples were taken for deter­
mination of needle weights, were dried to constant weight at 65°c'
Dried needles of each composite sample were ground to 40 mesh
in a stainless steel mill and stored in closed containers at
10°C
until analyzed.
IThe use of equipment brand names does not constitute recom­
mendation or endorsement by the U.S. Department of Agriculture.
Results and discussion
Stand and site properties
The Pacific silver fir sites studied varied greatly in their
key properties (Table 1). Elevation was highest (1300 m) at
site 11 and lowest (110 m) at site 16 where a perched water
table was evident during the winter. The sites represented
many soil types. The stands were 26 to 37 years old and
averaged 30.9 years. Associated vegetation was composed
of a variety of herbs, shrubs, and trees. The most common
shrub and tree species were, respectively, huckleberry
(Vaccinium spp.) and western hemlock.
Site productivity, expressed as site index at 50 years,
ranged from 11.8 m at site 8 to 24.1 m at site 14 (Table 2).
Terminal growth for the 3 years immediately before harvest
ranged from 29.0 to 73.3 cm/year. For all sites, the average
growth was 49.7 cm/year. This rate of growth compares
with 44.0 em/year for coastal western red cedar (Thuja
plicata Donn ex D. Don) (Radwan and Harrington 1986)
and with 55.6 em/year for western hemlock in the Cascades
RADWAN ET AL.
1431
TABLE 1. General attributes of Pacific silver fir sites
Site No.
Approximate
location
Elevation
(m)
1
2
North Mountain
North Mountain
3
4
Lookout
Suiattle Mountain
Morovitz Creek
5
6
Stand
age
(years)
Soil parent
material
880
37
Volcanic ash
Hemlock, huckleberry, dogwood, twinflower
1165
26
Volcanic ash
850
30
35
Sedimentary
Hemlock, huckleberry, dogwood, queen cup
beadlily
Hemlock, huckleberry, salal, deerfern
Hemlock, huckleberry, devil's dub,
870
Sulphur Creek
Foss River
1035
1025
Moon Lake
Bare Mountain
28
28
Volcanic ash
Basalt
Sedimentary
Huckleberry, beargrass
Huckleberry, boxwood, dogwood
Noble fir, Alaska cedar, huckleberry, alder
Hemlock, huckleberry, fool's huckleberry,
dogwood
Hemlock, Noble fir, Douglas-fir, beargrass
10
Hugo Lake
Ladd Mountain
11
Pinochle Creek
1300
32
35
28
30
12
Watch Mountain
1170
30
13
Mineral
1145
35
14
15
16
Humptulips
Quinault
Queets
640
775
110
28
33
31
Pumice and
volcanic ash
Pumice and
volcanic ash
Basalt
Sandstone
Glacial till
17
18
19
Raymond
Nelson Hill
Hyas Ridge
500
520
780
31
26
36
Basalt
Sandstone
Sandstone
29
salmonberry
Mountain hemlock, huckleberry
Hemlock, Douglas-fir, huckleberry,
twinflower
Basalt
Basalt
V olcanic ash
Volcanic ash
Volcanic ash
1140
1170
1220
1195
7
8
9
Associated plant species a
Hemlock, Douglas-fir, huckleberry, dogwood
Hemlock, Noble fir, huckleberry, twin flower,
beargrass
Huckleberry, Oregon oxaIis, foam flower
Hemlock, huckleberry, deerfern
Hemlock, red cedar, fool's huckleberry,
crabapple
Hemlock, salmonberry, fool's huckleberry
Hemlock, Douglas-fir, red cedar, salmonberry
Douglas-fir, huckleberry, salal
aHemlock, Tsuga heterophylla; huckleberry, Vaccinium spp.; dogwood, Comus canadensis; twin flower, Linnaea borealis; queen cup beadlily, Clinlonia
salal, Gaultheria shallon; deerfern, B/eehnum spicant; devil's club, Oplopanax horridum; salmonberry, Rubus spectabilis; mountain hemlock, Tsuga
merlensiana; Douglas-fir, Pseudotsuga menziesii; beargrass, Xerophyllum tenax; boxwood, Pachistima myrsinites; Noble fir, Abies procera; Alaska cedar,
Chamaecyparis nootkatensis; fool's huckleberry, Menziesia !erruginea; Oregon oxalis, Oxalis oregana; foam flower, Tiarella trifoliata; rcd cedar, Thuja plicata;
Crabapple, Pyrus !usea; alder, Alnus sinuata.
unij7ora;
(Radwan and DeBell 1980). Like other true firs, Pacific silver
fir trees exhibit rapid height growth for a fairly long period
of time following an initial stage of slow juvenile growth
(Harrington and Murray 1982).
The average needle number per centimetre of twig tissue
(23) and the average needle weight (8.1 mg) were much
higher than average values for western hemlock at
lOA needles and 2.0 mg per needle (Radwan and DeBell
1980). Also, the average needle area (both sides),
2
45.1 mm , appeared larger than areas of most associated
conifers. These needle properties could be among the
inherent characteristics of the species. Presumably, such
properties would represent special adaptations to take best
advantage of environmental conditions for survival and
growth at higher elevations.
Relationships between site and growth characteristics
As expected, site index was significantly (p :s; 0.01) cor­
related with terminal growth (r
0.89). Similar correlations
were obtained with western hemlock (Radwan and DeBell
1980) and with western red cedar (Radwan and Harrington
1986). Site index was also significantly (p :s; 0.01) related
to needle dry weight (r
0.54) and to number of needles
per centimetre of twig tissue (r
0.59). Other significant
(p :s; 0.01) relationships found were: needle dry weight vs.
needle area (r
0.55), terminal growth vs. needle dry
weight (r
0.57), site index vs. elevation (r
-0.77), and
terminal growth vs. elevation (r
0. 59). The last two
correlations show the strong negative relationship of eleva­
=
=
=
=
tion with site productivity and growth. The relationship
between site index and elevation is illustrated in Fig. 2.
Similar relationships are common among other western
conifers.
Foliar macronutrients
Foliar N concentrations ranged from 0.70 to 1. 06070
(Table 3). Unlike the information compiled by Minore
(1979), average total N (0.93%) was much lower than con­
centrations normally found in needles of other western coni­
fers, such as western hemlock (Radwan and DeBell 1980),
western red cedar (Radwan and Harrington 1986), and
Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco)
(Lavender and Carmichael 1966). Foliar N concentrations
of silver fir found in this study, however, were within the
species' range of values reported in the literature (Beaton
et al. 1965b; Gessel and Orians 1967; Cochran et al. 1986).
Similar low levels of N were also found in needles of other
true firs, such as California red fir (Abies magnifica
A. Murr.) and white fir (Abies concolor (Gord. and Glend.)
Lindl.) (R.F. Powers, USDA Forest Service, Redding CA,
personal communication). These low N levels and the fact
that the values in our study were mostly associated with good
tree growth and dark-green foliage suggest that silver fir (and
perhaps other Abies spp.) may have a low N requirement
resulting from a high efficiency of utilization of the element.
As with needle numbers mentioned before, a low N require­
ment may represent an ecological adaptation, since true firs
are commonly found at high elevations where soil temper­
CAN. J. FOR. RES. VOL. 19, 1989
1432
TABLE 2. Site index and growth characteristics of Pacific silver fir sites
Terminal growth
Site index
at 50 years
Needle no. per cm
of twig tissue
Needle
dry wt.
Site No.
(m)
per yeara
(cm)
8
12
11.8
12.6
12.9
29.0
40.3
33.3
33.4
24. 2
28.3
6. 3
6. 6
8.0
13.2
13.9
14.2
14. 2
14. 3
14. 3
33.3
44.3
31.8
21.4
20.0
28.2
22.9
7.9
7.0
8. 6
23.5
19.2
7.5
8. 6
8. 6
7
11
13
9
19
10
6
5
2
1
4
18
16
17
3
15
14
Avg.
SD
(mg)
Needle
area
2
(mm )
38.8
37. 0
44.4
37. 8
51.6
46. 6
45.2
15.4
16.1
39.3
31.3
46.7
46. 7
52.0
50.7
20.4
20.9
21.0
21.5
22.0
58.0
64. 0
62.7
50.0
73.3
22.3
22.7
24. 1
53.7
64.7
71. 3
20.2
18.0
21.6
23.6
7. 6
10.0
9. 5
42.6
41.8
62.0
43.4
59.2
47.2
17. 2
4.1
49. 7
13.1
23.0
4.5
8. 1
0. 9
45.1
7.0
23.2
19.7
15.9
19. 2
23.8
7. 8
9.2
7.8
8. 0
8.4
8.4
8.4
54.8
41.2
38. 2
46.8
40.2
39.0
aAverage terminal growth per year for the 3 years 1983, 1984, and 1985.
atures and nutrient mineralization are low. On the other
hand, the low N values of silver fir were well within known
deficiency levels of other western conifers. Silver fir and
perhaps other true firs, therefore, would likely benefit from
application of N fertilizer. Success of such treatment was
demonstrated more than 2 decades ago in a silver fir stand
in the northern Cascade Range of Washington (Gallagher
1964). More recently, good response to N fertilizer was also
reported in several red and white fir stands in the southern
Cascade Range (Powers 1981).
Sulphur concentrations ranged from 0.06 to 0.10070. Con­
centrations of < 0.10070 at most sites were similar to those
reported for interior silver fir and Douglas-fir (0.06-0.09070)
(Will and Youngberg 1979; Cochran et al. 1986) and for
coastal western red cedar (0.08-0.10070) (Radwan and
Harrington 1986). As with N, low levels of S indicate that
silver fir may have a low S requirement.
Ratios of S:N ranged from 0.08 to 0.11, and trends were
similar to those of S. These ratios were all higher than the
0.06 ratio normally found in healthy higher plants; they indi­
cate that there was sufficient S to balance the N present in
the formation of foliar protein.
Average concentrations of P (0.12070) and K (0.77070) were
lower than those reported by others (P
0.16-0.18070,
K
1.05-1.20070) (Beaton et al. 1965b; Will and Youngberg
1979). These concentrations, however, were within the low
end of the range of values found in other western conifers,
such as coastal Douglas-fir, western hemlock, and western
0.12-0.15070, K
0.50-0.90070) (Beaton
red cedar (P
et al. 1965b; Lavender and Carmichael 1966; Radwan and
==
==
==
==
DeBell 1980; Radwan and Harrington 1986). Many coastal
sites in western Washington are known to be especially low
in extractable P (Radwan and Shumway 1983).
Average levels of Ca (0.34070) and Mg (0.09070) were within
the ranges reported in foliage of silver fir and other true firs,
30
24
E
x
Q)
-0 c
Q)
18 12
•
-
U5
6
r
==
(n
==
-0.77
19)
0
0
500
1000
1500
Elevation (m)
FIG. 2. Relationship between site index and elevation .
Correlation coefficient (r) is significant at p ::s: 0.01.
as well as other western conifers, excluding western red cedar
0.20-0.70070, Mg
0.07-0.18070) (Beaton et al.
(Ca
1965b; Will and Youngberg 1979; Radwan and DeBell 1980;
==
==
R.F. Powers, personal communication). Average Mg values,
however, were barely above the critical value of 0.07070
proposed for red fir seedlings (Powers 1981); they were
especially low relative to Ca concentrations, resulting in high
Ca:Mg ratios. Previously, Krajina (1970) observed that soils
rich in both Mg and Ca were required for good growth of
silver fir.
Calcium levels found in this study were much lower than
the 1.65070 reported in foliage of old-growth silver fir by
Turner and Singer (1976). This very high Ca level was prob­
ably caused by analysis of a sample containing many old
needles. Silver fir is known to retain its needles for long
periods of time, and Ca is an immobile nutrient, which tends
to accumulate in older foliage.
RADWAN ET AL.
TABLE 3. Macronutrient concentrations
of Pacific silver fir
Site No.
N
P
Ca
K
(070) in foliage
Mg
S
0.94
0.11
0.69
0.45
0.10
0.09
0.12
0.10
0.98
0.70
0.27
0.51
0.Q7
0.11
0.08
0.10
0.96
0.99
0.81
0.96
0.94
0.94
0.99
0.09
0.12
0.13
0.13
0.13
0.13
0.12
0.74
0.70
0.79
0.88
0.79
0.80
0.79
0.39
0.28
0.30
0.32
0.26
0.32
0.29
0.10
0.08
0.07
0.12
0.10
0.10
0.08
0.09
0.08
0.08
0.09
0.08
0.10
0.08 11
12
0.76
0.13
0.80
0.24
0.09
0.08
0.97
13
14
15
16
17
0.96
1.02
1.06
0.70
0.13
0.14
0.69
0.74
0.28
0.36
0.10
0.11
0.10
0.09
0.11
0.11
0.09
0.60
0.80
0.93
0.37
OAI
OAO
0.11
0.10
0.09
0.08
0.08
0.06
4
5
6
7
8
9
10
18
19
1.06
0.94
0.84
0.10
0.10
0.12
0.73
0.74
0.69
OA2
0.31
0.36
0.08
0.08
0.10
0.08
0.08
0.Q7
Avg.
SD
0.93
0.09
0.12
0.01
0.77
0.09
0.34
0.07
0.09
0.01
0.08
0.01
TABLE 4. Concentrations (ppm) of micronutrients and
aluminum in foliage of Pacific silver fir
B
Cu
49
37
35
25
13
22
4
2
0.01
0.01
36
70
18
21
2
23
25
13
23
5
4
22
28
28
4
4
4
2
0.01
0.01
<0.01
0.01
Fe
1
2
1353
991
3
802
4
5
492
982
929
6
7
8
9
10
11
12
13
14
15
16
37
35
548
494
608
740
51
68
97
675
52
56
45
42
49
49
47
956
855
556
680
912
679
17
18
19
434
598
Avg.
SD
225
30
32
32
752
AI
Zn
Mn
Site No.
48
16
Mo
0.02
0.01
0.01
<0.01
234
248
173
160
192
99
305
272
231
93
33
22
19
19
28
20
36
19
30
25
25
25
20
14
4
2
0.01
<0.01
0.01
4
0.01
28
15
26
19
5
4
207
198
285
251
225
47
19
20
26
20
29
16
4
2
14
4
2
16
4
<0.01
0.01
0.01
0.01
<0.01
20
5
3
1
0.01
0.00
247
209
222
244
247
260
Foliar micronutrients and aluminum
Concentrations of the micronutrients and Al varied
among the 19 sites (Table 4). Very few comparisons can be
made between our values and those obtained by others
because the literature is extremely limited. Turner and Singer
(1976) reported concentration of one element in one sample
of old-growth trees (Mn = 1700 ppm), and Will and
Youngberg (1979) determined concentrations of four
elements in one sample of interior silver fir (Mn
825 ppm,
40 ppm, Zn
33 ppm, B
26 ppm).
Fe
Compared with important associated conifers, levels of
Zn, Fe, B, and Cu in silver fir foliage were within the ranges
=
=
=
=
26
•
24
0.91
0.96
1
2
3
1433
•
___ 22
E
x
(!)
"0 c:
(!)
U5
•
20
18
16
14
12
10
0.20
•
,
r
(n
•
__
__
__
==
0.75
=
__
0.30
19)
__
0.50
0040
Foliar Ca (%)
FIG. 3. Relationship between site index and foliar calcium
(r) is significant atp :5 0.01.
concentration. Correlation coefficient
of values reported for western hemlock, Douglas-fir, and
western red cedar (Zn = 15-35 ppm, Fe = 37-100 ppm,
13-57 ppm, Cu = 3-8 ppm) (Beaton et al. 1965a;
B
Radwan and DeBell 1980; Zobel and Hawk 1980; Radwan
and Harrington 1986). Concentrations of Mo (mostly
<0. 01 ppm) were much lower than those of other conifers,
especially red cedar (0.01-0.52 ppm) (Beaton et al. 1965a;
Radwan and DeBell 1980; Radwan and Harrington 1986).
Foliar levels of Al and Mn were higher than those found
12-86 ppm, Mn
69-383 ppm)
in western red cedar (Al
(Radwan and Harrington 1986) and lower than values for
550-1000 ppm, Mn
530western hemlock (AI
2000 ppm) (Beaton et al. 1965a; Radwan and DeBell 1980;
Zobel and Hawk 1980).
=
=
Foliar nutrients and site-stand characteristics relationships
Positive and negative correlations were found among
foliar nutrients and between the nutrients and important site
and stand characteristics (Table 5). As expected, N and S
were correlated (I'
0. 46); together with N, S is required
for synthesis of S-containing amino acids. The effect of Ca
on the availability of P was also manifested by the negative
-0.63).
correlation between the two elements (r
Correlation between foliar N and site index was not
significant. The importance of N to growth of silver fir,
however, was shown by correlations with terminal growth
(I'
0.48) and with needle area (I'
0.48). Similarly, Ca
was positively correlated with site index (r = 0. 75) (Fig. 3)
and with terminal growth (I' = 0.54), and was negatively
related to elevation (I'
-0.57). In contrast, P showed a
positive relationship with elevation (r = 0.81) and negative
- 0.86) and terminal
correlations with site index (I'
growth (r =
0.71). Clearly, these contrasting correlations
were related to the negative correlation between Ca and P
mentioned above.
Other correlations listed in Table 5 include those between
N and K (I' =
0.42), S and Mg (I'
0. 42), Fe and S
0.43), S and elevation
(I'
0.42), Zn and Fe (r
(I' = 0.54), Zn and terminal growth (I' =
0.42), and Zn
and needle weight (I'
-0.47). The importance of these
relationships is not immediately apparent.
Significant correlations between foliar nutrients and site­
stand characteristics suggest that some elements, such as P,
Ca, and N, may be useful as indicators to assess site quality
=
=
=
=
=
CAN.
1434
TABLE
J.
FOR. RES. VOL. 19, 1989
5. Significant correlation coefficients (r) and probabilities (p) among concentrations
of foliar nutrients, and between nutrient concentrations and site and growth characteristics
Correlation
r
p
Correlation
r
p
0/0 N vs. 0'/0 S
0/0 N VS. % K
0/0 P vs. % Ca
0/0 S vs. % Mg
ppm Fe vs. % S
ppm Zn vs. ppm Fe
Elevation vs. % Ca
Elevation VS. % P
Elevation vs. % S
0.46
-0.42
0.63
0.42
0.42
0.43
-0.57
0.81
0.54
0.023
0.035
0.002
0.035
0.035
0.032
0.005
0.001
0.001
Site index vs. % Ca
Site index vs. % P
Terminal growth vs. % N
Terminal growth vs. % P
Terminal growth vs. % Ca
Terminal growth vs. ppm Zn
Needle area vs. % N
0.75
-0.86
0.48
-0.71
0.54
-0.42
0.48
-0.47
0.001
0.001
0.018
0.001
0.008
0.036
0.018
0.022
for silver fir production (i.e., lower concentrations of P and
higher concentrations of Ca and N associated with better
sites). The same relationships also indicate that productivity
of silver fir may be improved by application of fertilizers,
especially those containing N and Ca; fertilization may be
particularly effective during the seedling stage when growth
is typically slow.
Acknowledgements
The authors thank the following organizations for their
cooperation in the study: ITT Rayonier, Inc., Murray
Pacific Corp., Weyerhaeuser Co., Olympic National Forest,
Gifford Pinchot National Forest, and Mt. Baker­
Snoqualmie National Forest. They also thank J.E. Wilcox,
D. W. Johnson, and S.R. Ray, Forestry Sciences
Laboratory, for their valuable assistance with the various
phases of the study.
BEATON, 1.D., BROWN, G., SPEER, R.C., MAcRAE, I., MCGHEE,
W.P.T., Moss, A., and KOSICK, R. 1965a. Concentration of
micronutrients in foliage of three coniferous tree species in British
Columbia. Soil Sci. Soc. Am. Proc. 29: 299-302.
BEATON, 1.D., Moss, A., M ACRA E, 1., KONKIN, 1.W., M CGHEE,
W.P.T., and KOSICK, R. 1965b. Observations on foliage
nutrient content of several coniferous tree species in British
Columbia. For. Chron. 41: 222-236.
BREMNER, 1.M., and MULVANEY, C.S. 1982. Nitrogen-total.
In Methods of soil analysis, part 2. Agronomy, 9: 595-624.
BUTTERS, B., and CHENERY, E.M. 1959. A rapid method for the
determination of total sulphur in soils and plants. Analyst
(London), 84: 239-245.
CHAPMAN, H. D. and PRATT, P.F. 1961. Methods of analysis for
soils, plants, and waters. Division of Agricultural Science,
University of California, Berkeley.
COCHRAN, P.H., LOPUSHINSKY, W., and MCCOLLEY, P.D. 1986.
,
Effect of operational fertilization on foliar nutrient content and
growth of young Douglas-fir and Pacific silver fir. USDA For.
Servo Res. Note PNW­445.
DIMOCK, E.l. 1958. Silvicultural characteristics of Pacific silver
fir. USDA Forest Service Pacific Northwest Forest and Range
Experiment Station, Portland, OR. Silvie. Ser. No. 4.
FOWELLS, H.A. (Compiler). 1965. Silvics of forest trees of the
United States. Agric. Handb. U.S. Dep. Agric. Sci. Educ. Adm.
No. 271. pp. 31-35.
FRANKLIN, J.F. 1982. The true fir resource. In Proceedings of the
Biology and Management of True Fir in the Pacific Northwest
Symposium, Seattle, WA, February 1981. Edited by C.D. Oliver
and R.M. Kenady. College of Forest Resources, University of
Washington, Seattle. pp. 1-6.
GALLAGER, L.U.
1964. A study of the effects of fertilization with
nitrogen and potassium on the growth and nutrition of Abies
Needle weight vs. ppm Zn
amabilis with associated greenhouse trials. M.S. thesis, Univer­
sity of Washington, Seattle.
GESSEL, S.P., and KLOCK, G.O.
1982. Mineral nutrition of true
fir. In Proceedings of the Biology and Management of True Fir
in the Pacific Northwest Symposium, Seattle, WA, February
1981. Edited by C.D. Oliver and R.M. Kenady. College of Forest
Resources, University of Washington, Seattle. pp. 77-83.
GESSEL, S.P., and ORIANS, G.H. 1967. Rodent damage to fer­
tilized Pacific silver fir in western Washington. Ecology, 48:
694-697.
HARRINGTGN, C.A., and MURRAY, M.D.
1982. Patterns of height
growth in western true firs. In Proceedings of the Biology and
Management of True Fir in the Pacific Northwest Symposium,
Seattle, WA, February 1981. Edited by C.D. Oliver and R.M.
Kenady. College of Forest Resources, University of Washington,
Seattle. pp. 209-214.
HEGYI, F., JELINEK, 1.J., VISZLAS, J., and CARPENTER, D.B.
1981. Site index equations and curves for the major tree species
in British Columbia. Forestry Inventory Report No.1. Ministry
of Forests, Inventory Branch, Victoria, B.C.
HORWITZ, W.
(Editor). 1980. Official methods of analysis of the
Association of Official Analytical Chemists. Association of Offi­
cial Analytical Chemists, Washington, DC.
1970. Ecology of forest trees in British Columbia.
In Ecology of Western North America. Vol. 2. University of
KRAJINA, V.l.
British Columbia, Vancouver. pp. 1-146.
LAVENDER, D.P., and CARMICHAEL, R.L.
1966. Effect of three
variables on mineral concentrations in Douglas- fir needles. For. Sci. 12: 441-446. MINGRE, D.
1979. Comparative autecological characteristics of
northwestern tree species. A literature review. USDA For. Serv.
Gen. Tech. Rep. PNW 8 7.
-
MURRAY, M.D., and TREAT, D.L.
1980. Pacific silver fir in the
coast range of southwestern Washington. Northwest Sci. 45:
119-120.
PACKEE, E.C., OLIVER,
C.D., and CRAWFORD, P.D. 1982.
Ecology of Pacific silver fir. In Proceedings of Biology and
Management of True Fir in the Pacific Northwest Symposium,
Seattle, WA, February 1981. Edited by C.D. Oliver and R.M.
Kenady. College of Forest Resources, University of Washington,
Seattle. pp. 19-34.
1976. Analytical methods of
atomic absorption spectrophotometry. Perkin-Elmer Corp.,
Norwalk, CT.
POWERS, R.F. 1981. Response of California true fir to fertiliza­
tion. In Forest Fertilization Conference Proceedings, Seattle,
PERKIN-ELMER CGRPORA TION.
WA, September 1979. Edited by S.P. Gessel, R.M. Kenady, and
W.A. Atkinson. College of Forest Resources, University of
Washington, Seattle. pp. 95-101.
RADWAN, M.A., and DEBELL, D.S.
1980. Site index, growth, and
foliar chemical composition relationships in western hemlock.
For. Sci. 26: 283-290.
RADWAN, M.A., and HARRINGTON, C.A.
1986. Foliar chemical
RADWAN ET AL.
concentrations, growth, and site productivity relations in western
red cedar. Can. J. For. Res. 16: 1069-1075.
RADWAN, M.A., and SHUMWAY, ].S. 1983. Soil nitrogen, sulfur,
and phosphorus in relation to growth response of western
hemlock to nitrogen fertilization. For. Sci. 29: 469-477.
SNEDECOR, G.W. 1961. Statistical methods applied to experiments
in agriculture and biology.
Ames, IA.
Iowa State University Press,
1435
T URNER, J., and SINGER, M.J. 1976. Nutrient distribution and
cycling in a sub-alpine coniferous forest ecosystem. ]. Appl.
Ecol. 13: 295-301.
WILL, G.M., and YOUNGBERG, C.T. 1979. Some foliage nutrient
levels in tree and brush species growing on pumice soils in cen­
tral Oregon. Northwest Sci. 53: 274-276.
ZOBEL, D.B., and HAWK, G.M. 1980. The environment of
Chamaecyparis lawsoniana. Am. MidI. Nat. 103: 280-297.
About this file: This file was created by scanning the printed publication. Some mistakes introduced by scanning may remain.