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WILDLIFE RELATIONSHIPS AND FOREST PLANNING
Steve Egeline
Biological Sciences Planner
Sierra National Forest
1130 11 0 11 Street
Fresno, California 93721
ABSTRACT
The complexity of the 11 Wildlife resource 11 is vastly
greater than any other resource area that the public or
private sector manages. To address this complexity,
knowledge of all the species has been collected in a single
reference (the Wildlife Habitat Relationships Program or
WHR) in such a manner that both analysis and decisions can
be intelligently made.
The issue in wildlife resources planning is not which
habitats are to be retained or even which are more important.
The primary issue is what distribution of relative abundances
among all the habitats will provide the most desirable mix of
wildlife within the demands for individual species and within
ecological considerations for wildlife diversity. The WHR
program has begun to enable land managers and biologists to
evaluate effects in a manner meaningful to address the issues
and concerns.
The question of 11 0ld growth 11 habitat, what it is and how
much might be desirable is discussed as an example of WHR
application in forest planning. Evidence from past descriptions
ecological theory and species adaption patterns obtained from
WHR data indicate that 11 0ld growth 11 consists of distinct habitats
and the relative abundances of these habitats are currently much
different than those existing prior to the influence of modern
man. The implications of this analysis to forest planning are
discussed.
KEYWORDS: Habitat, wildlife, land use planning, ecosystem, old
growth, mana_gement, diversity.
379
t_.',_,
I
A species' habitat consists of all the environmental features that provide the
food, cover, space and water necessary for population survival. This Huchinsonian
concept of habitat and ecological concept of species survival is the basis for the
wildlife-habitat relationships currently being developed (Thomas 1979, Salwasser 1980).
Pressure to ensure all species' survival has been manifested in leqal action by
various groups ultimately resulting in a plethora of laws and regulations (Salwasser
et al. 1980). These legislations and interpretations contain such words and phrases
as 11 • • • untrammeled by man's ... , ....... maintain viable populations ... , ....... diversity
of plant and animal communities ... ,
natural forest ... , 11 ... wildlife resour11
ce(s),11 and public .. l/. Few, if any, of these terms or concepts have been formally
defined in relationships to land management planning. In reality, the lack of formal,
written declaration of meaning for these terms is a problem only to lawyers and
planners since the legality of a forest land management plan is ultimately decided
by interpretation of the letter of these laws (i.e., the above phrases) in a courtroom. The writing of these and many other regulations on human behavior is an
attempt to legislate morality, in this case an ecological morality, and I believe it
can be readily demonstrated throughout history that moral legislation fails miserably
as soon as the moral atmosphere changes. Unfortunately, moral atmospheres change
more rapidly than laws can be promulgated and infinitely more rapidly than ecosystems
can react.
11
11
11
•••
These problems of moral, ecological, legal interplays impact directly on the
wildlife planner since the forest land management plan is written according to law
in response to public needs and demands (i.e., morals) within the context of the
forest's ecosystems.
In the absence of direct definitions, I have made some interpretations and
formulated conceptional frameworks that will be used in wildlife planning on the
Sierra National Forest. The first concept is that there is no "wildlife resource."
In reality, each species is a resource in itself, therefore, the term is 11 Wildlife
resources ... This may seem somewhat simplistic but it is not. A very great majority
of people concerned about Wildlife, .. both professional biologists and pressure
groups, do not have the conception of all wildlife species when they address the
wildlife resource. Generally, concern about the 11 Wildlife resource 11 and planning for
wildlife is done in the conceptual framework of a few, relatively important species
that are defined in each person's mind based on his experience, training, and overall
background (Warren et al. 1979). However, the legislated morality states "all
species'' and therefore it must be made abundantly clear to all people (individuals
and groups) that the wildlife resources being planned for are indeed each and every
species individually.
11
A second important concept, closely related to the first, is that there is no
11 public." If the wildlife planner can internalize the fact that every person is an
individual and possesses a unique conceptual framework it is less difficult to deal
with 11 public" issues and the wildlife resources. That is, there are 285 million
special interest groups which have some, but not all, concerns in common. An analogy
might be the "hunters ... This group has in common the concern for maintaining high
populations of huntable wildlife species. However, some of these hunters are more
concerned about bear than deer. This can lead to conflicting viewpoints about road
closure policies. Further, some bear hunters may use hounds in their sport and some
may not, again splitting the group.
Obviously this can be carried to the logical
extreme that I have stated above, but should nevertheless be a part of the wildlife
planners• concept of the public 11 if he is to deal with conflicting demands for species
and/or habitats.
1/ An excellent reference containing the legislation using these terms is "The Prin- ciple Laws Relating to Forest Service Activities, .. Agric. Handbook No. 453.
11
11
11
380
..
,•.
The third set of concepts, dealing with such terms as diversity, viability, natural forest, etc., are based in theoretical ecology but must be applied to practical
management. Many biologists are currently concerned over the interpretation of these
theoretical concepts as applied to the planning job, particularly in light of the
likely legal interpretation to be made of forest planning efforts. The remainder of
this paper deals with these concerns and is a result of my interpretation of the
laws, public issues and concepts of theoretical ecology and should not be construed
as nationwide Forest Service policy or direction.
I believe there will be some commonality in wildlife issues on a large majority
of western U.S. forests,if not throughout the nation. The major items concern
uncommon species (i.e., Endangered, Threatened, Rare, etc.), riparian habitat, the
effects of roads on wildlife, the influence of vegetation manipulation (i.e., timber
management, fuels management, etc.) on diversity and the question of how much "old
growth" is necessary for dependent species survival. These concerns, along with
nearly any others concerning wildlife and fish, are inextricably intertwined in the
fabric of the ecosystems. To illustrate this point let's consider the "old growth"
issue in the context of theoretical ecology and such things as "viability" and
"diversity."
The overall objective for any forest plan dealing with wildlife should be to
determine what proportion of each plant community containing what proportion of each
seral stage in what proximities and sizes will yield the best mix of wildlife species
and populations levels that meet the "overall multiple-use objectives" as constrained
by the ecosystems' abilities. This illustrates the complexity of wildlife planning
in that many things other than plant community and/or successional stage determine
"habitat."
In approaching this objective it is useful to apply some principles from gaming
theory. The first principle is that of uncertainty.
Particularly in the area of
ecosystems and habitats, it is generally true that there are fewer incorrect
decisions than correct decisions. This is often a function of knowledge deficiency,
that is, the only reason that there are fewer wrong answers than right answers is
that we know, through experience, what the wrong answers are but only know what the
right answers are through theory, which has rather nebulous confidence limits,
leading to a range of correct answers but no single correct answer. The second
precept of gaming that applies and is directly related is that of risk avoidance.
This principle states that as the amount at risk increases, the odds must increase
in favor of a correct decision or the bet will not be placed.
The application of these principles to wildlife planning should be apparent.
Decisions should be as conservative as possible and should be based on the values of
the habitats at risk. This is particularly true in decisions concerning vegetative
structure. It is relatively easy to correct a wrong decision concerning shrubs for
example, since it is relatively easy and quick to "grow" shrubs. However, if an
incorrect decision on the amount of old growth needed is made (on the low side) it
may be many hundreds of years before it can be corrected, which is obviously too
long with respect to the survival of dependent wildlife species.
The approach I've used in dealing with these problems is to assume that viable
populations of all wildlife species could be insured if the forest structure (plant
communities, seral stages, energy-flow patterns, etc.) mimics the conditions under
which the various wildlife species have evolved. The first problem is to determine
what the structure was.
In western forests it seems reasonable to assume that no plant community or seral
stage has been totally eliminated by modern man. It should therefore be reasonable to
381
assume that all plant communities and seral stages now present on a forest were also
present during the evolution of wildlife species. Thus, our first criteria is to insure in the forest planning process that no extant plant community or seral stage
thereof can potentially be eliminated by prescribed activities. While this seems an
intuitively obvious decision I believe that each step in the planning process needs to
be made visible and apparent, and this in fact is called for by law.
Our second criteria deals with the question of the relative abundances of the
various plant communities and seral stages prior to the influence of modern man.
Various studies have pointed out that many plant communities were subject to periodic
fires (Kilgore 1973; Thomas 1979; Thompson and Taylor 1979) which maintained them in
a particular structural arrangement. And that structure differs considerably from
the climatic climax structure. In that modern man began effectively controlling
these low-intensity fires around the turn of the century, it should also be obvious
that few if any people now alive are personally familiar with the pre- modern-man
forest. This is apparent when old growth forest is considered. This habitat is
generally thought of as a large tree, multistoried, dense canopy forest with small
openings, a high degree of decadence, and many logs and other debris on the ground.
Photographic evidence (Gibbens and Heady 1964; Progulske and Sowell 1974) and the
written descriptions by many early explorers including John Muir, John C. Fremont, and
various land surveyors does not paint the same picture. The forests described prior
to fire control were indeed characterized by large trees, but there the similarity
ends. The "natural forest•• old growth also had a relatively sparse canopy closure,
few but young and vigorous shrubs and little accumulation of litter or logs.
11
11
While the evidence for a larqe-tree, open-canopy forest is fairly apparent, the
sample size is fairly limited. Also, in that we are dealing with a "high risk"
decision, it would seem desirable to gather as much evidence as possible (i.e.,
increase the odds of a correct decision) prior to committing a particular ecosystem
to an essentially irreversible decision. The Western Sierra Wildlife Habitat Relationships (Verner et al. 1980) has arranged data for each wildlife species in such a
manner that the quality (optimum, suitable, etc.) of a particular plant community and
seral stage can be determined with some degree of confidence for each species. By
examining these data, certain patterns of adaptation can be discerned. Figure 1
illustrates the number of wildlife species that find optimum or suitable living
conditions in each of the 70 defined plant community/seral stage combinations. Figure
2 is simply.a refinement of Fig. 1 by considering only those species that show strong
reaction to canopy closure and tree size. This second group excludes the reptiles,
amphibians, and bird and mammal species that depend on such things as water, cliffs,
caves, etc. Figure 3 shows how many of these tree/canopy dependent species are most
influenced by large trees with closed canopies, large trees with open canopies and
smaller trees with similar canopy closures. It is readily apparent that large trees
with open canopies (less than 70 percent canopy closure) are an important structure
to more species than any other arrangement.
The ecological theories that apply are those of speciation and partitioning.
The more abundant and stable a resource is, the more advantageous it is to become
specialized on a portion of that resource since specialization reduces resource competition; that is, the resource can be more finely divided without consequence to the
species. This leads to the conclusion, supported by the mentioned written and photographic evidence, that "old growth 11 habitat in fact was primarily the large-tree,
open-canopy type with the closed-canopy type being secondary in abundance and stability. However, if one then examines the particular species that require the closed
old growth type9 one can see that these species are of equal concern with those
adapted to open old growth (Table 1).
The conclusion to be drawn is that there are at least two 11 0ld growth 11 habitats;
one being large trees with less than 70 percent canopy closure, single layer tree
382
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0
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J:
Q)
...
....~
Q)
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Figure 1.
Percentage of all wildlife species that find optimum or suitable breeding, feeding or resting
habitat in each of the plant community/seral stage combinations. Plant communities include:
annual grasses (ANNGRS), blue oak shrub (BLOS), digger pine (DIGP), chaparral (CHAP), ponderosa
pine (PONP), California black oak-white fir (BKOW), mixed conifer (MCON),'jeffrey pine (JEFP),
red fir (REDF) and lodgepole pine (LOOP). Seral stages are: (1) = grass-forb stage, (2) =
shrub-seedling-sapling stage, (3) =pole-medium tree stage and (4) = large tree stage. Canopy
cover classes are: (A) = from 0 to 39 percent canopy cover, (B) = from 40 to 69 percent canopy
cover and (C) = 70 percent or more canopy cover.
V-1
00
-1:>-
8?.
~
0
CD
....c
Figure 2.
Habitat use by those wildlife species which are strongly influenced by canopy
closure and tree size. The height of bars indicates the percent of these species
which find optimum or suitable breeding, feeding or resting habitat in each of the
plant community/seral stage combinations. For explanation of symbols, refer to
Fig. 1.
~
00
CJ1
a..
Q)
I.-
0
Q)
c
....,
.....0
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rn
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Q)
.....0
~
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....,
I
DIGP
c(
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:t:
tRill
a.
~
PONP
BKOW
MCON
JEFP
REDF
11213 4 31413 4 1l2J3 434 11213 4 31413 4 11213 4 31413 4 11213 4 31413 ~
I A ale
lA ale
lA a
lA ale
lA ale
The optimum habitats of those wildlife species whose occurrences seem to be
associated with tree size and canopy closure. For explanation of symbols,
refer to Fig. 1.
BLOS
11213 4 314 11213 4 3T4l3l4
lA ale
lA a
Figure 3.
II~
en
l§
~
_....
LOOP
Table 1.
Species of the western Sierras whose distribution is influenced
by tree size and canopy closure in old growth stands.
Closed Old Growth
Open Old Growth
Yuma myotis
Wolverine (?)
Little brown myotis
Fisher
Ruby-crowned kinglet
Marten
Mountain chickadee
Black bear
Steller 1 s jay
Northern flying squirrel
(Myotis yumanensis)
( Gu lo luscus)
(Myotis lucifugus)
(Martes pennanti)
(Regulus calendula)
(Martes americana)
(Pa:t'US gambeli)
(Ursus americanus)
(Cyanocitta stelleri)
(Glaucomys sabrinus)
Olive-side flycatcher
Western qray squirrel
(Nuttallornis borealis)
(Sciurus griseus)
Western wood pewee
Douglas
(Contopus sordidulus)
1
squirrel
(Tamiasciurus douglasi)
Black-backed three-toed woodpecker
Golden-crowned kinglet
White-headed woodpecker
Brown creeper
Hairy woodpecker
Red-breasted nuthatch
Williamson 1 s sapsucker
Chestnut-backed chickadee
Yellow-bellied sapsucker
Pileated woodpecker
Common flicker
Great gray owl
(Picoides a:t'cticus)
(Regulus satrapa)
(Picoides albolarvatus)
(Certhia familiaris)
(Picoides villosus)
(Sitta canadensis)
(Sphyrapicus thyroideus)
(Parus rufescens)
(Sphyrapicus varius)
(Dryocopus pileatus)
(Colaptes auratus)
(Strix nebulosa)
Band-tailed pigeon
(Columba fasciata)
Blue grouse
(Dendragapus obscurus)
American kestrel
(Falco sparverius)
Goshawk
(Accipiter gentilis)
Pine grosbeak
(Pinicola enucleator)
Cassin s finch
1
(Carpodacus cassinii)
Evening grosbeak
(Hesperiphona vespertina)
Western tanager
(Piranga ludoviciana)
Hermit warbler
(Dendroica occidentalis)
386
canopy, few logs and debris,and sparse, vigorous shrubs; the second characterized by
large trees with dense, multilayered canopy and high fuel loadings. The relative
proportions of these two types on any given National Forest can best be determined by
close consultation with fuels management specialists to determine which areas on the
forest were likely not subject to periodic fire due to local topography and soil/moisture regimes. A general conclusion is that the closed old growth likely was associated
with high elevation, north slope, riparian areas and that open old growth was associated with hotter, drier sites.
A secondary conclusion to be drawn is that the ponderosa pine and mixed conifer
plant communities are not in the same proportion as existed previous to modern man's
influence, since many timber stands identified as mixed conifer are, in fact, large
ponderosa pine trees with an understory of fire sensitive species such as whjte fir
and incense cedar. This is further supported by the adaptive patterns shown in Fig.
2. More species find optimum habitat in the ponderosa pine type than in mixed conifer.
The management conclusions and recommendations to be made concerning "old
growth" might be to manage north-facing mixed conifer communities within one-quarter
mile of water as closed old growth in the amount determined as existed naturally
(within that l/4 mile band) by consultation with fuels management specialists. The
open old growth might best be managed on poorer timber-producing sites in ponderosa
pine and mixed conifer since these sites generally have poor stocking and exhibit
the open crown closure characteristic. These open old growth stands can be maintained by periodic individual tree selection cutting and regular prescribed fire.
This will likely result in substantial reduction of closed old growth habitat
acreages but it has been argued here that the "natural forest" probably contained
substantially fewer acres of this habitat than now exist on most national forests and
that those dependent species should be adapted to that situation.
A final note on "old growth" is that species requiring this habitat structure
tend to also require rather large patches, therefore old growth of either type should
be managed in a minimum patch size of 300 acres and an average of about 600 acres.
It should also be apparent that blanket statements of ''leave x percent of each timber
compartment, capability area, etc., in old growth" are entirely inadequate since the
land varies in its ability to produce large trees and the "natural" placement of these
habitats should be determined by terrain, surrounding vegetation, etc.
Using these analytical techniques it is also possible to display "diversity."
The ecological characteristics of diversity such as richness, equitibility and pattern
can all be displayed if acres are substituted for number of species and the size statistics of mean, mode, range and standard deviation are included in the histogram for
each community/seral stage. Figure 4 is an example of how this might be done. By
displaying components of diversity in this manner and relating the species to the
components through a wildlife-habitat relationships concept, it should be relatively
easy to address many wildlife concerns by alternative. This can be done by displaying
a Figure 2, Figure 3 and Figure 4 for the "natural forest," the "existing forest" and
the "future forest" under each of the alternatives and including a wildlife species
list for each of the community/seral stage combinations. In this manner, anyone with
a concern for an individual species or group of species can determine the effects of
the various alternatives. Also, these characteristics of habitat richness, equability
and size are measurable, enabling numeric goals to be set and monitored.
A final concept
habitat feature that
species for planning
increase as a result
in planning is to consider only those species that depend on a
will be reduced by management activities. That is, eliminate
consideration that depend on habitat structures that will
of some management activity. Also, only consider those plant
387
I.
r
I
I.
I
communities that will be impacted and finally only consider those management activities that have major, widespread impact in relation to the habitat or species being
impacted. This process should result in the 11 bettering of the odds 11 in those 11 high
risk 11 areas and investigation into which decisions are incorrect rather than which are
correct. Management direction can then be written to constrain 11 Wrong 11 decisions
rather than to attempt creating 11 right 11 decisions. The optimum decision in land
management is that which maintains the maximum range of future options within the
constraints of current needs.
10000
Fig. 4.
An example of the acreages (numeric goals) of each plant
community/seral stage combination that might be desirable
for wildlife habitat--see text.
Acknowledgments
Conversations with Jared Verner, Hal Salwasser, Bruce Marcot, Ed Toth, and Hugh
Black, Jr. have served to stimulate the thoughts presented in this paper.
Literature Cited
Boyce, Stephen G. and Noel D. Cost.
1978. Forest diversity - new concepts and applications.
Research Paper SE-194.
388
USDA Forest Service
Gibbens, R. P. and H. F. Heady.
1964. The influence of man on the vegetation of Yosemite Valley, California.
Agric. Exp. Stn. Manual 36. 44 p.
Kilgore, Bruce M.
1973. The ecological role of fire in Sierran conifer forests, its application to
national park management. ~Journal of Quarternary Research 3(3).
Lyon, Jack L., Hewlett S. Crawford, Eugene Cenhai, Richard L. Fredricksen, Richard
F. Harlow, Louis J. Mete, and Henry A. Pearson.
1978. Effects of fire on fauna, a state-of-knowledge review. USDA Forest
Service Gen. Tech. Report W0-6.
Progulske, Donald R. and Richard H. Sowell.
1974. Yellow ore, yellow hair, yellow pine; a photographic study of a century
of forest ecology. Bulletin 616, Agric. Exp. Stn., South Dakota State Univ.,
Brookings, S. D.
Salwasser, Hal, John C. Capp, Hugh Black, Jr., and Janet Hurley.
1980. The California wildlife habitat relationships program: an overview.
~ Proc. of the workshop on management of western forests and grasslands for
nongame birds. USDA Forest Service Gen. Tech. Report, lntermt. For. and Range
Exp. Stn., Ogden, Utah.
Thomas, Jack Ward, technical editor.
1979. Wildlife habitats in managed forests, the Blue Mountains of Oreqon and
Washington. 512 p. Agriculture Handbook No. 553, U.S. Government Printing
Office, Washington, D. C.
Thompson, Rita R. and Alan R. Taylor.
1979. The environmental significance of fire.
~Biology
Digest.
Verner, Jared.
1980. Bird communities of mixed-conifer forests of the Sierra Nevada. In
Proc. of the workshop on management of western forests and grasslands for
nongame birds. USDA Forest Service Gen. Tech. Report, lntermt. For. and Range
Exp. Stn., Ogden, Utah.
Warren, Charles, Marshall Allen, and James W. Hackner.
1979. Conceptual frameworks and the philosophical foundations of general living
systems theory. Behavioral Science 24:296-310.
389
t ' ''