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Density as a Misleading Indicator of Habitat Quality

Article  in  Journal of Wildlife Management · October 1983

DOI: 10.2307/3808148

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Density as a Misleading Indicator of Habitat Quality
Author(s): B. Van Horne
Source: The Journal of Wildlife Management, Vol. 47, No. 4 (Oct., 1983), pp. 893-901
Published by: Wiley on behalf of the Wildlife Society
Stable URL: http://www.jstor.org/stable/3808148
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DENSITY AS A MISLEADING INDICATOR OF HABITAT QUALITY

B. VAN HORNE, Department of Biology, University of New Mexico, Albuquerque, NM 87131

Abstract: Current methods of evaluating wildlife habitat for management purposes can be arranged
hierarchy of increasing generality. The most general level is evaluation of wildlife habitat for entire
munities on the basis of inferences drawn from vegetational structure. At the base of the hierarchy the
resolution studies, upon which accuracy at the higher hierarchical levels depends, usually assume that h
quality for a species is positively correlated with the density of the species. If habitat quality for a wil
species is a measure of the importance of habitat type in maintaining a particular species, habitat q
should be defined in terms of the survival and production characteristics, as well as the density, of the
occupying that habitat. Situations in which habitat quality thus defined is not expected to be posi
correlated with density are described, along with the species and environmental characteristics that are
likely to produce these situations. Examples drawn from the literature in which density and habitat qu
are not positively correlated are described. The positive correlation of density with habitat quality in s
instances cannot be assumed without supporting demographic data.
J. WILDL. MANAGE. 47(4):893-901

decoupled. In such cases, management

The foundation of any wildlife habitat
policies based directly on species abun-
management plan is the ability to assess
habitat quality accurately. Without dance
this may be misleading and these errors
may be amplified when management ap-
key ingredient, the effort put into care-
fully prepared objectives and elegant cat-
proaches are restricted to the higher levels
egorizations of habitat types is largely
of the hierarchy.
wasted. Yet biologists often dwell on ob-This paper is dedicated to the late O.
jectives and categories while treating C. Wallmo, who was always eager to dis-
cuss ideas and whose refusal to be any-
lightly the assumptions implicit in their
assessments of habitat quality. For thingin- but completely honest in evaluating
stance, they seldom question the assump-his own ideas, objectives, and research
tion that the density of a species in a ideas,
hab- as well as those of others, set an
itat is a direct measure of the quality of
example for us to follow.
that habitat. Perhaps this is because any
METHODS OF HABITAT
more accurate investigation of habitat
EVALUATION
quality to truly reflect the importance of
Habitat assessment procedures can be
that habitat in maintaining wildlife species
populations must be intensive, often atvisualized
the in a 3-level hierarchy of in-
expense of the broader informationcreasing
base generality in which the accuracy
that could be achieved by simple surveys.
of predictions at 1 level is dependent on
Such surveys are a particularly common accuracy at the next lower level (Fig. 1).
means of evaluating nongame wildlife
The lowest level is the assessment of the
habitat. habitat relationships of individual species
The objectives of this paper are to pro-
at a particular site. Accuracy at this level
is dependent upon an intimate under-
vide some examples of situations in which
this correlation does not hold, and to make
standing of the demography of the species
predictions regarding species and envi- and of the factors influencing population
ronmental types for which the density- levels through their influences on survival
habitat quality relationship is likely to beproduction, although such analyses are
and

J. Wildl. Manage. 47(4):1983 893

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894 HABITAT QUALITY . Van Horne

LEVEL 3
often abandoned in favor of simple esti-
mation of total density. Biases of the dif- INDIRECT EVALUATION OF
ferent censusing techniques are a prob- HABITAT QUALITY FOR
lem, particularly when these are habitat z
O A WILDLIFE COMMUNITY
dependent, as are many of the bird cen-
susing techniques (Emlen 1971) and most LEVEL 2

mammal censusing techniques where o INDIRECT EVALUATION OF

U)
home range size varies among habitats. At w HABITAT QUALITY FOR A
the next higher level, the species-habitat ar
SINGLE SPECIES USING
relationships are extrapolated to sites not (9
actually sampled. The success of this in- z INFERENCES FROM LEVEL I
ference is dependent upon the correct LEVEL I
identification of the important factors in-
Ilr DIRECT EVALUATION OF
fluencing density in the higher resolution
w HABITAT QUALITY FOR A
studies. At the highest levels of the hier-
SINGLE SPECIES USING
archy, the extrapolated information is used
ON-SITE DATA
to put together a habitat quality assess-
ment for an entire wildlife community. Fig. 1. Hierarchical description of habitat quality ass
Generally, in evaluating the effects of ments.
management options on the wildlife com-
munity, species interrelationships such as
competition and predation are ignored and
the community assessment is based solely the generalist wildlife species common to
on the aggregation of individual species disturbed habitats and may ignore the
assessments. sensitive species with greater habitat spec-
Over the last decade there has been ificity. Second, habitat diversity and wild-
considerable pressure to developlife rapid
species diversity are not always posi-
means of habitat-quality assessment,tively
suchcorrelated; this depends on the ratio
of base
that the higher-resolution levels at the generalist to specialist wildlife species
of the hierarchy (Fig. 1, levels 1 in
and 2)area being managed and the spe-
the
are skipped altogether. For instance,cific
in 1requirements of those species.
approach maximizing species diversity is
Assessment of a range of habitat types
assumed to be the primary objective of presence or absence of wildlife
for the
management and this diversityspecies is as-is a procedure at the 2nd level of
sumed to be directly correlated withresolution.
hab- The general objective in this
itat diversity (Asherin et al. 1979).case
There
is to manage lands so that sufficient
are several problems associated with thistypes are retained to allow for rep-
habitat
approach. First, maximum diversity resentation of all species while maximiz-
achieved in the limited areas beingingman-diversity within this constraint
aged (a diversity) may not produce(Thomas
max- 1979). Determination of the
presence or absence of species is usually
imum diversity on a larger scale (0 diver-
sity) (Samson and Knopf 1982), because
based on the available literature; there is
some wildlife species, such as old-growth
no explicit treatment of the densities re-
quired
specialists, are not adapted to areas of high to avoid extinction (minimum vi-
habitat diversity. Maximizing plant able
com-populations) or of home range size
and no
munity diversity on a local scale selects forevaluation of habitats on the basis

J. Wildl. Manage. 47(4):1983

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 HABITAT QUALITY * Van Horne 895

of wildlife species densities or the relative

sity as an indicator of habitat quality than
favorability of the occupied habitats. do assessments at higher levels of gener-
Another example of a procedure at thisality, it has been suggested that the ac-
2nd level of resolution is the U.S. Fish and curacy of any habitat rating technique,
Wildlife Service's Habitat Evaluation Pro- such as HEP, should be tested by com-
cedures (HEP) (Flood et al. 1977). Thisparing habitat ratings to the observed rel-
procedure relies on assessment of habitatative abundances of a variety of wildlife
requirements of individual species taken
species (Whelan et al. 1979).
from the literature, followed by assess- Thus, the assumed positive correlation
ment of habitat types based on the ability of a species' abundance with habitat qual-
of each type to provide for these require- ity underlies most methods used for as-
ments. The 2 major problems with this sessing habitat quality and is explicit for
approach are that our knowledge of spe- the species-specific level of resolution. It
cies requirements is often poor and syn- is therefore the basis for a broad range of
ergistic effects among resources are ig- management decisions regarding wildlife
nored. Thus, a habitat that provided cover communities.
but no food would still get a positive value The assumed relationship often breaks
rating, even though the species might not down under intensive study. One reason
be able to exist in that habitat. Likewise, that it may break down, particularly in
the sum of 2 "good" resource attributes northern climates, is that habitat use in
might well be greater than their separatewinter is critical, whereas most censuses
values. and surveys are taken in warmer months.
All 3 of the above methods allow for
For northern deer (Odocoileus spp.), the
rapid habitat assessment without direct
availability of winter range may contrib-
censusing of wildlife species. They are thus
ute disproportionately to carrying capac-
based on untested inferences about which
ity (and thus survival and reproductive
species "ought" to occur in each habitat
patterns) (Wallmo et al. 1977). Identifi-
type and are not suited to management cation of habitat quality on the basis of
for viable population levels. summer densities would thus be mislead-
ing; retention of the summer habitat type
ASSUMPTIONS OF HABITAT
would not serve to maintain the popula-
EVALUATION
tion if the winter range was destroyed.
The assessment of individual species-
A 2nd reason for a breakdown in the
habitat relationships is the lowest leveldensity-habitat
of quality assumption is that
the hierarchy; these data are critical to there
the may be multi-annual variability in
success of the HEP type of analyses. The local population densities that reflects
usual assumption at this level is that small-scale
the variability in the food source,
local density of a species is positively cor-
in predator populations, or in abiotic en-
vironmental factors. Densities may thus
related with habitat quality. Often a range
of habitat variables is measured and cor- reflect conditions in the recent past or
related with species density. Multivariatetemporary present, rather than long-term
procedures are used to reduce the number habitat quality. For instance, site tenacity
in breeding passerines can produce local
of variables and to aid in interpretation of
results (e.g., Carey 1980, Maurer et al.densities that reflect past, rather than cur-
rent, habitat quality (e.g., Hild6n 1965,
1980). Although this type of habitat as-
Rotenberry and Wiens 1978).
sessment depends more explicitly on den-

J. Wildl. Manage. 47(4):1983

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896 HABITAT QUALITY * Van Horne

Third, social interactions may prevent vival probability, and mean expectation
subdominant animals from entering what of future offspring, for residents in 1 area
is actually the high-quality habitat, while as compared to other areas. More precise-
at the same time suppressing reproduction ly,
in the high-quality habitat. The surplus
individuals may then collect in habitat
"sinks," where densities may fluctuate
widely (Lidicker 1975). Animals in the low
quality sinks survive and/or reproduce
poorly. Thus, in a good year, the source nxj(la,,Bx, + Px,)
nxj (1/a3)
population may produce a large excess of
juveniles that will emigrate and build up
to high densities in the sinks. Because the
juveniles are subdominant, there is no so-
cial interaction factor to prevent high , n(lBx,/ + P))
densities in the sink habitats, which is in
contrast to the adult-dominated high-
quality or source habitats. Densities in the
lower-quality habitat may thus actually be where Qi is the relative quality of habitat
greater at times than in the high-quality i for the species, Bx is the fecundity of an
habitat. A similar scenario is embodied in
x-year-old and I, the probability that the
the theoretical model of habitat occupan-offspring will survive to a, the Ist age of
cy developed by Fretwell and Lucas breeding. Px is the probability of surviving
(1969). In this model, the movement of from age x to age x + 1, n is the number
individuals into poor habitat is a reflectionof individuals in each of the i habitats
of individual fitness maximization. Ac- being compared, and a is the area that
cording to the model, the per-individual
includes all sampled individuals in the ith
probability of success for unestablishedhabitat. The areas must encompass the
immigrants may be higher in low-quality home ranges of the individuals included.
Conceptually, this is a measure of mean
than in high-quality habitat, because high
individual "fitness" per unit area. "Fit-
densities in the high-quality habitat pro-
mote a high probability of failure toness"
re- is used here in a management rather
produce successfully and a high mortalitythan an evolutionary context; it describes
rate among the unestablished immigrants. a mean group characteristic in 1 habitat
Thus, it may be individually advanta- as compared to other habitats, rather than
geous for them to settle in the lower-qual-
comparing 1 individual of a population to
ity habitat. other individuals of the population. The
measure of habitat quality thus has com-
DEFINITION OF HABITAT
ponents of density, offspring production,
QUALITY
and survival. High density alone does not
Fitness of an individual animal (Fisher
infer quality habitat. To give an extreme
1930) is a relative measure that increasesexample, one could imagine a habitat in
with increasing survival probability andwhich all animals were immigrants and
increasing offspring production. I proposenone emigrated or reproduced. The qual-
that habitat quality be defined as the ity of the habitat would be zero. If either
product of density, mean individual sur-
individual survival probability or number

J. Wildl. Manage. 47(4):1983

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 HABITAT QUALITY * Van Horne 897

z demographic study of single species in a

0
variety of habitats. In this way one can
separate low-quality habitats, which may
0- contain largely immigrants that are un-
likely to survive or reproduce well, from
higher-quality habitats, containing a low-
0. er density of animals but in which densi-
a-
ties are more stable, reproductive output
of the population is dependable, and the
population is more likely to persist in poor
:D
or "crunch" years. Where such intensive
demographic study is impractical, den-
sity-based estimates could be greatly im-
proved through attention to immigration
patterns, to adult survival, and to the pro-
duction of juveniles that survive to repro-
t LOW HIGH duce.
HABITAT QUALITY The actual parameters used in equation
1 will be means for a certain time period,
Fig. 2. Change in minimum viable population size with in-
creasing habitat quality. commonly a year. An accurate assessment
of habitat quality requires the calculation
of a grand mean and variance over several
such time periods. The number of time
of offspring produced is zero, thenperiods
the required for a useful measure of
habitat quality will be greater for highly
habitat is making no contribution toward
maintaining populations of the speciesunpredictable
and habitats.
its quality is zero. There are some problems inherent in
the use of this habitat quality measure.
Given this definition of habitat quality,
the minimum viable population sizeThe will
areas encompassed by habitat patches
a, may in some cases influence survival
be greater in low-quality than in high-
quality habitat, because low survivaland production characteristics, particular-
and
production rates in low-quality habitat
ly for wide-ranging species. This will re-
mean that a higher density is neededsultto
in lower Q,'s for smaller patches con-
ensure persistence of the species intaining
that habitat equivalent in quality to that
habitat (Fig. 2). of the larger patches. Area of the patches
To measure habitat quality, one must
considered is thus an implicit variable in-
determine the mean production andfluencing
sur- Qi. Calculating Qj's for large or
vival characteristics of each age-class for
and similar-sized patches will remove the
the number of resident individuals in each
area effect. Calculating Q1's for different-
age-class in each habitat. Such a deter- sized patches with similar habitat char-
mination will be impractical for most acteristics will make the area effect ex-
studies. The above formula is thus pre-plicit. Also, home ranges may encompass
sented to clarify the definition of habitat several patches of what we perceive as dif-
quality and provide an ideal standard. This ferent habitat types, and the delineation
measure of habitat quality may be ap- of areas for which favorability is to be de-
proximated sufficiently through intensivetermined must thus be somewhat arbi-

J. Wildl. Manage. 47(4):1983

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898 HABITAT QUALITY * Van Horne

trary. Further, the patches used by a because there was no breeding by young
species may be from widely separated of the year.
areas, as for migratory birds. In such cases, Two different habitat types were distin-
it may be useful to make separate assess- guished for both adults and juveniles by
ments of summer and winter range, and discriminant function analysis: those char-
of the degree to which different habitat acterized by high densities or low densi-
types in each of these ranges contribute to ties of the age-class. Individual animals
mean relative fitness. whose home ranges encompassed high-
I have defined habitat quality in terms density adult habitat had a significantly
of a single species. The habitat quality forhigher probability of surviving over the
a wildlife community is the sum of habitatwinter period than those whose home
qualities for species members, as modified ranges encompassed low-density adult
by the effects of species interaction. I havehabitat, whether these animals were adults
discussed the problems of simply equatingor juveniles. Thus, high-quality habitat
habitat quality with diversity. Although it could be distinguished by the adult habi-
has been asserted (Cringan et al. 1979) thattat discriminant function and appeared to
more community-level research is needed be positively correlated with overwinter
as input to the development of habitatsurvival for both age-groups. The opposite
management plans, a valid assessment ofwas true of the juvenile discriminant
the effects of habitat manipulation at thefunction, for which the habitat character-
community level is dependent upon theized by high densities of juveniles con-
accuracy of assessments at the individualferred lower overwinter survival proba-
species level. In most cases, our under- bility. Thus, high-density adult habitat was
standing of individual species-habitat re-of high quality, while high-density juve-
lationships is still rudimentary. nile habitat was of low quality.
These quality inferences were corrob-
EXAMPLES
orated by the observation that adult male
Several examples of situations can
weights
be on the grid containing mostly
considered in which habitat qualityhigh-density
and adult habitat were signifi-
species density are not positively correlat-
cantly higher than those on the other trap-
ed, because of the influence of social ping grids and the population density on
dominance factors. In my own studies of this grid was relatively stable. However,
a series of populations of Peromyscusin 1979, the last year of the study, total
maniculatus in spruce (Picea spp.) and densities on those grids containing mostly
hemlock (Tsuga spp.) stands of differentlow-quality habitat exceeded those on the
seral stages in southeast Alaska (Van Hornegrid containing mostly high-quality hab-
1982), the populations occurred at a highitat. This was due to irruptions of juveniles
latitude (550N) where there was no breed- that consisted largely of immigrants that
ing by young of the year and the domi-were probably forced into the lower-qual-
nant adults were clearly separable from ity habitat. Additional evidence for the
the subdominant juveniles on the basis of importance of intraspecific dominance in-
weight, pelage, and trapping history. Be- teractions in these populations came from
cause of forced emigration, reproductivebreeding inhibition in high-density pop-
success in different habitats was difficult ulations and from the observation that
to estimate. Overwinter survival, how- subdominant juvenile diets were of lower
ever, was a critical component of fitnessquality when these animals were found in

J. Wildl. Manage. 47(4):1983

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 HABITAT QUALITY * Van Horne 899

high-density populations. Thus, the den- ferred habitats when densities were high.
sities measured in 1979 would have been These species included the wren (Trog-
a completely misleading indicator of over- lodytes troglodytes), the chiffchaff (Phyl-
all habitat quality. loscopus collybita), the great tit (Parus
Other studies of small mammal popu- major), the yellowhammer (Emberiza ci-
lations have reported similar patterns. trinella), and the Eurasian kestrel (Falco
Kock et al. (1969), for example, found the tinnunculus). For these species, density
highest densities of lemmings (Lemmus would be a reasonably good measure of
lemmus) during a population "peak" far- habitat quality in years of low-overall den-
thest from the optimum habitat as defined sity, but would be a misleading indicator
by food availability. Animals in the lower-in years of high-overall density.
quality habitat tended to be smaller andWhen breeding birds are territorial and
were probably younger subdominants. favorable habitat is limited, a surplus of
States (1976) reported that subdomi- adults of breeding age ("floaters") may
nant yellow-pine chipmunks (Tamias accumulate in poor habitat where either
amoenus) accumulated in marginal hab- no breeding takes place or where breed-
itat where their survival probability was ing attempts are largely unsuccessful.
relatively low. A large component of these Thus, a group with low current "fitness"
marginal populations consisted of immi- may be found in moderate densities in
grants. Thus, the marginal areas appeared poor habitat. This phenomenon has been
to be acting as dispersal sinks for animalsreported for great tits (Krebs 1971), the
forced out of the central areas, and den- Santa Cruz Island scrub jay (Aphelocoma
sity in the range of habitats investigated coerulescens) (Atwood 1980), and the
was not correlated with habitat quality. Australian magpie (Gymnorhina tibecen)
In an in-depth radio-tracking study, (Carrick 1963).
Schantz (1981) found similar numbers of
PREDICTIONS
red foxes (Vulpes vulpes) in mineral soil
and peat soil habitats. He was able to Problems with assuming density to be a
identify the mineral soil habitat as being measure of habitat quality are thus found
of higher quality despite the similarity over
in a wide range of taxa. We are left
density, as it contained a higher propor- with several important questions. To what
tion of reproducing adults. extent can we extrapolate these findings
Similar observations have been made for to other species? How general is this lack
breeding passerines. Fretwell (1969) re- of close relationship of density to habitat
ported that there was "no positive corre- quality? Where do we expect to find den-
lation between density and suitability" forsity and habitat quality to be decoupled?
breeding field sparrows (Spizella pusilla) I suggest that this phenomenon might be
where suitability was measured in termsfound in association with 3 main environ-
of breeding success; densities were higher mental types (Table 1). The Ist is highly
in an area where breeding success wasseasonal habitat in which different habitat
lower. O'Connor (1981) summarized data types may be preferred at different sea-
for a number of migrant and non-migrant sons, such that the density-habitat quality
bird species in Great Britain. The speciesrelationship cannot be inferred from sur-
showed a pattern of filling only a certain, veys or censuses taken during only 1 sea-
presumably preferred, habitat when den- son. The real high-quality habitat in this
sities were low, but filled the less pre- situation would be that which in some way

J. Wildl. Manage. 47(4):1983

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900 HABITAT QUALITY * Van Horne

Table 1. Factors that increase the probability that density

will not be positively correlated with habitat quality.
of species in those cases where habitat-
quality ratings are based on actual survey
Environmental characteristics Species characteristics or census data. This is because generalists
Seasonal habitat Social dominance
interactions
are relatively easy, to survey and are more
likely than specialists not only to occupy
Temporal unpredict- High reproductive a wide range of habitats, but to be found
ability capacity
Patchiness Generalist
in high densities in at least some habitat
types and to have a high reproductive ca-
pacity. The 3 species characteristics are
more closely associated with small, than
with large, body size.
was most critical for successful individualIt is likely that for rare species, density
survival and reproduction. A 2nd environ-may remain a reasonably good indicator
mental attribute is unpredictability overof habitat quality if seasonal changes in
time. This would allow for opportunistic
habitat use are taken into account and if
density increases in low-quality habitat, orhabitat is not patchy. If the habitat is
overflow into lower-quality habitats dur- patchy, the presence of a rare species in a
ing periods of high production and highgiven patch will have a larger stochastic
overall density. Third, habitat must beelement than the presence of a common
patchy on a scale allowing for migrationspecies in a habitat patch, because of the
between patches if environmental unpre- susceptibility of rare species to local ex-
dictability is to produce wide densitytinctions (e.g., Hanski 1982).
changes in the resident animals of 1 hab-
itat type relative to other habitat types.MANAGEMENT IMPLICATIONS
High densities in low-quality habitat could Management plans that depend only on
not be observed if there was no source habitat characteristics to infer habitat
pool in nearby high-quality habitat. quality contain a large amount of guess-
I would predict 3 main species charac-
work, both with regard to viable popula-
teristics to be associated with the habitat
tion levels and with regard to predicata-
quality-density decoupling (Table 1).
bility of species densities on the basis of
First, the species should have a social pat-habitat characteristics. Such plans depend
tern of dominance interactions where it isheavily on the correct identification of fa-
found in stable populations in high-qual-vorable habitat for the wildlife species
ity habitat. This type of dominance socialbeing managed. Intensive multi-annual
interaction is common to a wide range ofdemographic study of a single species over
vertebrates. Its demographic effects are
the range of habitats being measured is
most pronounced in those animals with aneeded to interpret the broader surveys.
2nd species attribute, high reproductive Without attention to demography, even
capacity. This high reproductive capacitymulti-annual surveys or censuses will not
can allow "sink" populations to reach high
necessarily be sufficient to distinguish
densities when the environment becomes "source" and "sink" habitats. Manage-
temporarily favorable. Third, this decou- ment plans adopted on the basis of a
pling should be most characteristic of hab- species survey or census taken during only
itat generalists. This is particularly impor- 1 year, or on the basis of measured habitat
tant as such generalists may be used as characteristics coupled with inadequate
indicators of habitat quality for a variety knowledge of the factors actually deter-

J. Wildl. Manage. 47(4):1983

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 HABITAT QUALITY * Van Horne 901

mining habitat quality, are particularly 1969. Notes on behaviour and food supply of
unsatisfactory. lemmings (Lemmus lemmus, L.) during a peak
density in southern Norway, 1966/67. Zeits. flir
Thus, we cannot afford to ignore the Tierpsychol. 26:609-622.
processes that produce the densities we KREBS, J. R. 1971. Territory and breeding density
observe, or attempts to maintain target in the great tit Parus major L. Ecology 52:2-22.
LIDICKER, W. Z., JR. 1975. The role of dispersal in
densities by retaining areas of specified the demography of small mammals. Pages 103-
habitat types will founder. We need to be 128 in F. B. Golley, K. Petusewicz, and L. Rysz-
much more careful in identifying high- kowski, eds. Small mammals: their productivity
and population dynamics. Cambridge Univ.
quality or critical habitat and not assume Press, New York, N.Y.
simple density-habitat quality relation- MAURER, B. A., L. B. MCARTHUR, AND R. C.
WHITMORE. 1980. Habitat associations of birds
ships without the demographic data to
breeding in clearcut deciduous forests in West
support them. Virginia. Pages 167-172 in D. E. Capen, ed. The
use of multivariate statistics in studies of wildlife
LITERATURE CITED habitat. U.S. Dep. Agric., For Serv. Gen. Tech.
Rep. RM-87.
ASHERIN, D. A., H. L. SHORT, AND J. E. ROELLE. O'CONNOR, R. J. 1981. Habitat correlates of bird
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J. Wildl. Manage. 47(4):1983

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