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Studies in Avian Biology No. 16:41-45,
1994.
INFLUENCE OF SITE QUALITY AND STAND DENSITY ON
GOSHAWK HABITAT IN SOUTHWESTERN FORESTS
RICHARD L. BASSETT, DOUGLAS A. BOYCE, JR., M. HILDEGARD REISER
RUSSELL T. GRAHAM, AND RICHARD T. REYNOLDS
Abstract. Current management guidelines for the Northern Goshawk (Accipitergentilis)in the Southwest call for a mosaic habitat consisting of approximately 10 percent of the forest area in grass-forb/
shrubs, 10 percent in 2.5-12.7 cm trees, 20 percent in 12.7-30.5 cm trees, 20 percent in 30.545.7
cm trees, 20 percent in 45.7-6 1.O cm trees, and 20 percent in 6 1.O cm and greater trees. This habitat
mosaic was conceived as convenient categories to describe a generally balanced, ecologically sustainable, forest ecosystem. In reality, however, the vegetative structural percentages vary. We describe
how differences in site quality and stand density affect vegetative structural stage percentage and forest
age.
Key Words: Accipitergentilis;forest regulation; Northern goshawk; site quality; stand density, SDI;
vegetative structural stages.
A goshawk scientific committee developed
recommendations for managing the Northern
Goshawk (Accipitergentilis) in the southwestern
United States (Reynolds et al. 1992). These recommendations focused on developing and maintaining forest conditions to provide habitat for
sustaining goshawks and their key prey species.
The recommendations defined size, location,
stand structure, woody debris, and soil condition
requirements for nest, post-fledging family, and
foraging areas. Stand structure included such
properties as the proportion and distribution of
six different diameter classesor vegetative structural stages(VSS), and number of large (>46 cm)
trees, snags, and down logs per ha.
Reynolds et al. (1992) recommended a mosaic
of vegetative structural stages interspersed
throughout the post-fledging family and foraging
areas in small, less than 1.7 ha patches to form
a balanced, ecologically sustainable, uneven-aged
forest. Vegetative structural stageis a generalized
description of forest structure and age based on
the majority of trees in a specific diameter class
within the forest (Table 1). The mosaic included
patches ranging from grass-forb/shrub to old forests, with a high priority on sustaining as much
as 40 percent of the area in mature and old forests. The recommended average proportion was
about 10 percent of the goshawk management
area (14,820 ha) in grass-forb/shrubs, 10 percent
in 2.5-l 2.7 cm (1-5 in.) trees, 20 percent in 12.730.5 cm (5-12 in) trees, 20 percent in 30.545.7
cm (12-l 8 in) trees, 20 percent in 45.7-6 1.O cm
(18-24 in) trees, and 20 percent in 61.0 cm (24
in) and greater trees.
In this paper, we examine how differences in
site quality and stand density influence VSS percentage and forest age. This paper also describes
the impacts of varying VSS proportions within
goshawk management areas to goshawk habitat
and their key prey species.
FOREST DEVELOPMENT
Reynolds et al.% (1992) recommended VSS
and forest mosaic can be approached by traditional, even-aged area regulation. Area regulation consists of dividing the forested post-fledging family and foraging areas into as many I 1.7
ha patches as there are years in the expected life
of a forest (forest age) and regenerating an equal
percentage of the forest each entry period (Smith
1986). The entry period could vary from 10 to
20 years depending upon existing forest conditions.
Three basic principles apply to area regulation.
First, structural stagesbecome important to sustain a forest over time, even where the desired
condition is to have large, old trees. Second, new
trees must be established at regular intervals to
sustain the desired structural stages in a forest
through time. Third, forests are dynamic. Trees
regenerate and grow at different rates and die at
different ages, resulting in a forest that is constantly changing over time (Oliver and Larson
1990).
Four variables that affect VSS distribution and
forest age are: (1) length of stand establishment
period, (2) site quality, (3) stand density, and (4)
tree longevity.
STAND ESTABLISHMENT
Length of stand establishment varies by species, regeneration method (natural or planting),
amount and kind of forest floor disturbance, and
climatic variation. The observed stand establishment period (or years in VSS 1) for ponderosa
pine (Pinus ponderma) forests can range from 15
years on a highly productive site to 30 years on
41
42
STUDIES
IN AVIAN
BIOLOGY
NO. 16
TABLE 1. VEGETATNESTRUCWRALSTAGFLS
AND pine seedlings and 4 large reserve trees (trees >
46 cm dbh) were simulated over a 320-year period at SDIs of 113 and 157. Four reserve trees
Vegewere allowed to die (snag creation) at stand age
tative
30, and thereafter thinning from below, to the
stnlcDiameters’
tura1
specified SDI, was allowed at 20-year intervals,
stage
Forestdescription
(cm)
starting at stand age 40 years.
1
O-2.5
Grass-forb/Shmb
Generally, seeding/sapling and young trees
Opening
have a faster diameter growth rate than mature
2
2.5-12.7
Seedling/Sapling
and old trees (Table 2). The time it took for a
12.7-30.5
3
Young
tree to move through one VSS ranged from 11
4
30.545.7
Mid-aged
to 59 years for 25% maximum SDI, and from
5
Mature
45.7-61.0
11 to 95 years for 35% maximum SD1 depending
>61.0+
6
Old
on the site quality and VSS. Also, it takes longer
’ Tree diametermeasuredat I .4 m abovegroundlevel.
to grow through a stage on the low productivity
sites than on high productivity sites; the excepa poor site. Generally, south- to west-facing, drytion was VSS 3 on average and high sites.
er slopes with shallow soils require the longest
seedling establishment period, north-facing, more
mesic sites with deeper soils require the least STAND DENSITY
time. Under similar topographic and soil conStand density influences tree diameter growth.
ditions, sites where annual precipitation is usuOn sites with the same site quality, tree diameter
ally less require more time than when moisture
growth will vary with different management inis more plentiful.
tensities. For example, the number of years in
VSS 4, 5, and 6 are longer for the higher stand
SITE QUALITY
density (35% SDI) than for the lower stand denSite quality influences tree growth and varies
sity (25% SDI). The growth simulator model
greatly in the Southwest. Site quality denotes the
showed no differences in number ofyears for VSS
relative productivity of a site for a particular tree
2. The seedling establishment period for low (30
species (Ford-Robertson 197 1). Factors that inyears), average (20 years), and high (15 years)
fluence site quality include soil characteristics,
site quality was assumed to be equal for the two
mineral composition, slope, aspect, microclidensities.
mate, and tree species (Daniel et al. 1979). Site
Stand density and site quality also influence
index is useful to help quantify site quality, and
VSS percentage and the time required to achieve
refers to the average height of dominant and cothe desired forest structure for goshawks and their
dominant trees in a stand at an arbitrarily chosen prey species (Tables 3 and 4). One desired forest
age. Minor (1964) developed site index curves
structure condition is to maintain 40% of the
for ponderosa pine in northern Arizona, meagoshawk post-fledging family and foraging areas
suring age at 1.4 meters above ground level using in VSS 5 and 6 to sustain moderate to high popa chosen base age of 100 years. For example, a
ulations of key prey species. These older age
site index of 70 (a tree 21.3 m [70 ft] tall at 100
classesmaintain the most speciesat an abundant
years of age at 1.4 m above ground level) is conpopulation level (10 of 12 species found in ponsidered about average for ponderosa pine in the
derosa pine forests: i.e., woodpeckers, chipSouthwest. Thus, the length of time required for
munks, tassel-eared squirrels) (Reynolds et al.
trees in each VSS is a function of site quality and
1992). Since forest stands reach VSS 5 and 6 from
stand density (Table 2).
30 to 70 years earlier in stands with lower density
A forest growth simulator model (Edminster
(25% SDI), the desired forest structure could be
et al. 199 1) was used to project diameter growth
maintained for a longer period of time. Olderper decade (Table 2) for two key stand densities:
aged stands (> 200 years) are also more frequent(1) Stand density index (SDI) 113 (25% of maxly used as goshawk nest sites. Because nest sites
imum SD1 450 for ponderosa pine); and (2) 157
have a higher density of large trees, these areas
SD1 (35% of maximum SDI). The first level is
should be managed for even higher density stands
considered to be the onset of competition be(43% SDI).
Actual VSS percentage varies from the rectween trees, whereas the second is the lower limit
of full site occupancy (Long and Daniel 1990).
ommended lo- 1O-20-20-20-20 (Reynolds et al.
Stand density index is the number of trees at an
1992). For example, the VSS percentage for a
low quality site with 25% SD1 is 12-9- 18- 17-23average stand density of 25.4 cm (10 in) (Daniel
et al. 1979, Lilliholm et al., this volume).
2 1 and for a high quality site with 25% SDI, VSS
During modeling, the growth of 500 ponderosa
is 9-6-22-l 9-22-22 (Table 3).
THEIR DWETERS
SITE QUALITY
AND
GOSHAWK
et al.
HABITAT--Bassett
43
DIA~~ETER
GROWTHANDNUMBEROFYEARSIN EACHVEGETATNESTR~~~~RALSTAGE
TABLE 2. ESTIMATED
(VSS) FORDIFFERENTQUALJTYPONDEROSAPINESITESANDSTAND
DENSITIE~INTHESOIJTHWEST
Stand density’
Dieter
Site quality’
@owth per decade
(cm)
25% Max SD1
Approxima;S;ars
35% Max SD1
in each
25% Max SD1
35% Max SD1
30
20
15
30
20
15
1
LOW
Average
High
2
LOW
Average
High
4.32
6.86
9.40
4.32
6.86
9.40
24
15
11
24
15
11
3
LOW
Average
High
3.68
4.83
4.15
3.05
4.32
4.24
48
37
37
58
41
42
4
Low
Average
High
3.48
4.06
4.83
2.79
3.25
3.63
44
38
32
55
47
42
5
Low
Average
High
2.59
3.63
3.99
1.60
2.44
2.91
6
LOW
Average
High
1.52
2.08
2.52
1.27
1.70
2.08
59
42
38
-3
95
63
51
-3
Seedling
estabishment
period
Seedling
establishment
period
’ Site quality (Minor 1964). Low = 50 SI (SI = site index - dominant tree height at 100 years), Average = 70 SI, High = 90 SI.
2 Stand density index (SDI) is the number of trees of average stand diameter of 25.4 cm. 25% max SD1 = onset of competition;
lower limit of full site occupancy.
6 depends on the selected forest age.
’ Years in VSS
35% max SD1 =
ponderosa pine is closer to 200 years or less
(Pearson 1950, White 1985, Covington and
Moore 1991). Life expectancy for Engelmann
spruce (Picea engelmannii) ranges from 250-450
years (Alexander and Shepperd 1990). The life
expectancy of the typical tree would be more
appropriate to set targets for sustaining forests
then the age of the oldest tree. Tree species and
TREE LONGEVITY
Tree longevity influences forest life expectancy
and forest age required to achieve desired forest
structure. The lifespan of trees varies within and
between species. For example, the oldest known
living ponderosa pine tree in the Southwest was
found to be 742 years old (Swetnam and Brown
1992), whereas the average life expectancy of most
DLNVIETER
GROWTH,YEARSIN VEGETATIVE
STRUCXWRAL
TABLE 3. ESTIMATED
STAGE,ACCUMULATED
AGE,
AVERAGE,ANDHIGHQUALITYSITES
ANDPER~ENTOFLANDXAPEINEACHVSSFORPONDEROSAPINEONLOW,
MAX
SDI)’
WHERE STAND
DENSITY
IS 113 SD1 (25%
Vegetative structure stages
Site quality
vss
1
vss 3
vss 4
vss 5
VSS 6’
Diameter growth/decade
Years (Act-years)’
% in VSS
0
30 (30)
12
4.32
24 (54)
9
3.68
48 (102)
18
3.48
44 (146)
17
2.59
59 (205)
23
1.52
55 (260)
21
Average(70 SI)
Diameter growth/decade
Years (Acc-years)3
% in VSS
0
20 (20)
10
6.86
15 (35)
8
4.83
37 (72)
18
4.06
38 (110)
19
3.63
42 (152)
21
2.08
48 (200)
24
High (90 SI)
Diameter growth/decade
Years (Act-years)”
% in VSS
1: (15)
9
l?$)
6
3;‘7;63)
22
3;.8:95)
19
3;‘;l933)
22
3;$70)
22
Low
(50
SI)
vss 2
’
Maximum stand density index for ponderosa pine is 450.
1 Number of years in VSS 6 is determined by selecting a growth period that is approximately
3 Number of years in VSS and accumulated years.
20% of forest age.
STUDIES
44
TABLE 4.
ESTIMATED
DWTER
GROWTH,
IN AVIAN
YEARS
IN
VEGETATIVE
OF LANDSCAPE IN EACH VSS FOR PONDEROSA
AND PERCENT
WHERE STAND DENSITY IS 157 SD1 (35% MAX SDI)l
NO. 16
BIOLOGY
PINE
STAGE,
ACCUMULATED AGE,
AVERAGE, AND HIGH QUALITY SITES
STRUCIWZAL
ON
Low,
Vegetative structural stages
Site quality
vss
1
vss 2
vss 3
vss 4
vss 5
vss 6’
Diameter growth/decade
Years (Act-year@
% in VSS
30 (30)
9
4.32
24 (54)
7
3.05
58 (112)
18
2.79
55 (167)
17
1.60
95 (262)
29
1.27
65 (327)
20
Average (70 SI)
Diameter growth/decade
Years (Acc-years)3
% in VSS
0
20(20)
9
6.86
15 (35)
7
4.32
41 (76)
18
3.25
47 (123)
20
2.44
63 (186)
27
1.70
44 (230)
19
High (90 SI)
Diameter growth/decade
Years (Acc-years)3
% in VSS
0
15(15)
7
9.40
11 (26)
6
4.24
42 (68)
21
3.63
42(110)
21
2.97
51 (161)
25
2.08
39 (200)
20
Low
(50 SI)
0
’ Maximum stand density index for ponderosa pine is 450.
2 Numberof Yean in VSS 6 is determined by selecting a growth period that is approximately 20% of forest age.
3 Number
of years in VSS and accumulated years.
longevity must be considered when considering
an older forest age.
By decreasing or increasing forest age, VSS
percentage is changed. For example, if the forest
age on the low productive site was lowered from
327 to 250 years, the VSS percentage would
change from 9-7-18-17-29-20
to 12-10-23-2233-0 (Table 4). VSS 6 may not be achievable if
a 327-year forest age is not ecologically sustainable. This should not influence goshawks, given
the average life expectancy of ponderosa pine
and that sites with low productivity (SI I 50)
account for only 2.3% (N = 4 national forests)
of ponderosa pine stands in the Southwest (USDA
1993). Planning for forest ageslessthan 200 years,
however, could negatively impact goshawks and
their prey populations. Regardless of site quality
or stand density index (25% or 35%), forests less
than 200 years will not provide for the older
classes(VSS 5 and 6) (Tables 3 and 4). Suitable
goshawk nesting habitat is commonly composed
of older trees (>200 years) in the VSS 5 and 6.
Prey species like the Red-naped Sapsucker
(Sphyrupicus nuchalis) and Williamson’s Sapsucker (Sphyrapicus thyroideus) would lose 66%
of the forest structure conditions that maintain
high populations (Reynolds et al. 1992). Forests
without VSS 5 and 6 would also not provide the
large snags that are used by other nesting-cavity
prey species.
TABLE 5.
STRUCTURAL
APPROXIMATE
PERCENT
RV EACH
VEGETATIVE
SUSTAINING
GOSHAWK
HABITAT
Not all structural stagesare equally important
for the goshawk and its prey species,but all structural stages are equally important for a forest to
become established and to sustain itself from the
grass-forb/shrub stage (VSS 1) with seedlings
through the old forest stage (VSS 6). The traditional area even-aged method of regulating a forest can be applied successfully to sustain a forest
with the mosaic of VSS that will meet the habitat
needs of the goshawk and its key prey species.
Of 12 goshawk prey species found in ponderosa pine forests, openings (VSS 1) are of no importance to 5 prey species(i.e., sapsuckers, tasseleared squirrel), and important to 1 prey species
(cottontail) for maintaining high populations. For
only 1 prey species (tassel-eared squirrel) found
in the ponderosa pine forests younger-aged forests (VSS 3) are important, and only when larger,
STAGE
(VSS)
AND
FOREST
AGE
THAT
CAN
BE EXPECTEDTO OCCUR IN GOSHAWK POST-FLEDGINGFAMILY AND FORAGING AREAS OF AVERAGE SITE QUALM
FOR PONDEROSAPINE FOREST TYPE AND MANAGEMENT INTENSITIES(REYNOLDS ET AL. 1992)
Percent in each vegetative structural stage
Management
intensity
management*
Minimal
Moderate
Intensive
No
I
Forest age
vss
1
10
9
10
10
vss 2
vss 3
10
13
10
8
80
20
19
18
vss 4
0
17
17
17
vss 5
vss 6
0
20
20
21
0
21
24
26
Unthinned ponderosa pine stand at Fort Valley Experimental Forest (Reynolds et al. 1992). An untbinned
yield model, never grew beyond VSS 3 with a 200-year forest age (Ronco et al. 1985, Edminster et al. 1991).
stand, using GENGYM
(ye.-)
200
233
204
194
growth and
SITE QUALITY
AND
GOSHAWK
older trees are available for nesting and seed
sources (Reynolds et al. 1992).
Under varying management options, VSS percentages never attain lo- 1O-20-20-20-20. The
youngest forest age (shorter time span) occurs
under intensive management; oldest forest age
(longest time span) occurs under minimal management (Table 5). Minimal management level
is characterized by trees that are significantly
competing with one another. When management
intensities are at moderate and intensive levels,
trees develop without significant competition.
Without management, however, unthinned ponderosa pine stands of average site quality are
unlikely to grow beyond the young forest structural stage (VSS 3) even after 200 years (Ronco
et al. 1985, Edminster et al. 1991).
To provide the desired forest conditions of
large old trees over 40% of the goshawk management area and small forest openings for prey
species and tree regeneration, entry periods for
management activities (i.e., harvesting, fire, etc.)
would need to be about every 20 years for the
moderate level. An expected level of management intensity (stand density) must be determined prior to establishing the desired VSS proportions and forest age.
The recommended 10-10-20-20-20-20
VSS
percentage is now being considered as a hardand-fast rule by those implementing and reviewing timber sale projects. However, the 10-10-2020-20-20 distribution was intended to describe
approximate percentages of each VSS throughout the post-fledging family and foraging areas
to sustain suitable goshawk habitat (Reynolds et
al. 1992). The achievable VSS percentage should
be determined by considering existing local factors that inlluence forest establishment and
growth, expected management intensity, and tree
longevity.
ACKNOWLEDGMENTS
We would like to thank the following reviewers:W.
M. Block, R. M. Jeffers,M. Johnson,M. J. Larson, R.
J. Lilieholm, C. 0. Minor, and M. L. Morrison.
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COVMGTON, W. W., AND M. M. MOORE. 199 1.
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HABITAT--Bassett
et al.
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