
Description
This is a layer representing predicted nesting habitat
for the Mexican Spotted Owl (Strix occidentalis lucida), as well
as a measure of nesting habitat quality, across the assessment area.
The
layer was built using a combination of simple rules based on a literature
review, and Mahalanobis distance based on habitat conditions at known
nest sites. The layer has a resolution of 90m (0.8 ha or 2 acres),
and values within the layer range from zero to 5. Higher values indicate
a higher degree of habitat suitability.
Purpose
This data layer was created as part of the ForestERA project
to support landscape-scale forest restoration planning efforts by
a broad group of stakeholders including federal and state agencies, academic
institutions, and non-governmental entities. These data are intended
for regional analyses over spatial extents on the order of tens to
hundreds of thousands of acres, and were not developed for use at
finer
spatial scales, although they may be useful for some applications
at finer scales.
Development
We chose two modeling approaches to optimize
the use of available information on Mexican Spotted Owl. First, because
previous
studies documented characteristics of nesting habitat for these species,
we used rules developed from the literature to identify the extent
of potential nesting habitat on the landscape. This involved a search
of the literature to identify a set of variables related to nesting
habitat and a range of possible threshold values that might be used
to identify areas most likely to support nesting pairs of these raptors.
Second, we employed the Mahalanobis distance statistic (hereafter M-distance)
to generate a dataset that could be used as a surrogate for habitat “suitability” or “preference”.
Georeferenced locations of 132 nest sites for Mexican Spotted Owl were
obtained from the United States Forest Service Rocky Mountain Research
Station and the United States Fish and Wildlife Service for use in
the modeling effort.
To identify potential nesting habitat, we used
publications that described habitat characteristics around Mexican
Spotted Owl nest sites (Grubb et al.
1997, May & Gutierrez 2002; Seamens & Gutiérrez 1995), breeding
season roost sites (Ganey & Balda 1994; Young et al. 1998, Zwank et al.
1994) and current management guidelines (USFWS 1995) to choose a range of threshold
values (Prather et al., in review). Since owsl are tied primarily to pine-oak
and mixed-conifer habitats, we initially created a “preferred vegetation” layer
with these two vegetation types identified as “habitat”. While
areas of pure ponderosa pine are not typically used by owls for nesting (USFWS
1995) we included areas identified as ponderosa pine in immediate proximity
(1 pixel or 90m) of pine-oak and mixed conifer to include ecotonal areas between
habitat types that may have been misclassified using the ETM imagery. Finally,
we included areas of pure ponderosa pine on steep slopes (>8o) because owls
have some affinity for areas with high slope (Ganey & Balda 1989a; Zwank
et al. 1994), and in our vegetation mapping we found that it was difficult
to distinguish pure pine from mixed-conifer or pine-oak on slopes >8o using
ETM imagery (ForestERA unpublished data). We then eliminated patches of habitat
smaller than 40 ha because these patches would likely be too small to be suitable
for nesting (USFWS 1995).
After delineation of potential nesting habitat, we
used M-distance and habitat characteristics at Mexican Spotted Owl Nest sites
to develop an index of habitat
suitability within potential nesting habitat. M-distance has been used frequently
in mathematics and physics to compare the similarity of datasets, but is
rarely used in ecology (e.g., Clark et al. 1993; Corsi et al. 1999; Knick
and Dyer
1997). M-distance is a multivariate statistic that provides a measure of
dissimilarity between two multivariate datasets (Farber & Kadmon
2003). In this effort, one dataset is the mean vector of habitat characteristics
at known Mexican
Spotted Owl nest sites while the other dataset is the range of conditions
across the entire landscape (Prather et al., in review). The "distance” for
any location on the landscape represents the dissimilarity between conditions
at that location and the mean habitat conditions at the known nest sites.
This value is scaled in multivariate space using the covariance between
the measured
habitat conditions at the locations used by the species (Clark et al. 1993).
We used dominant overstory vegetation, canopy cover, basal area, tree density,
slope, and the sine and cosine derivatives of aspect as predictor variables
for our M-distance modeling effort. We determined values for each of these
variables at Mexican Spotted Owl nest sites by extracting the values from
the GIS raster layers using the “zonal statistics” function in
ArcGIS Spatial Analyst (ESRI Corporation). We overlaid the nest locations
with the
raster layers of the predictive variables and determined the value of each
raster layer in the pixel corresponding to the nest location. Analysis of
characteristics of the 90m (0.81ha) pixel in which each nest fell was deemed
appropriate since
other studies have shown significant patterns between nest site selection
by large raptors and habitat characteristics at scales of approximately 0.5
-
1ha (e.g., McComb et al. 2002). For purposes of developing the M-distance
model we used 2/3 (82) of the nest sites as training data and the remaining
1/3 (41)
of the nest sites were withheld for accuracy assessment.
The final habitat
layer represents a combination of our two modeling efforts. We classified
this layer into 5 categories for purposes of display, analysis,
and dissemination. The zero category represents areas not predicted to
be suitable for nesting habitat using our simple rules. The remaining
4 categories
represent
areas of increasing similarity to the mean habitat characteristics at Mexican
Spotted Owl nest sites based on M-distance. Fifty percent of nest sites
would be expected to fall into category 4, an additional 25% into category
3, an
additional 15% into category 2, and the final 10% into category 1.
Accuracy
Assessment
To test the effectiveness of using threshold values within
predictor variables as a means of identifying potential
habitat, we used Chi-squared goodness of fit tests (Zar 1999) to determine
whether the potential habitat models did significantly better at predicting
locations of Mexican Spotted Owl nests than chance alone. For this
analysis we used the entire set of nest sites (n = 132). This analysis
suggested that significantly more (111 or 84%) nest sites fell within
the predicted extent of potential nesting habitat than would be expected
by chance alone (v = 2, X2 = 92.2, P < 0.0001).
To test
whether the M-distance statistic was effective at identifying the
locations of nest sites, we used Chi-squared goodness of fit tests
to determine whether
the M-distance statistic accurately predicted the number of nests within
the test datasets that fell within the portion of the predicted habitat
in each
of the four categories identified above. For these analyses we used the 1/3
of the nests (n = 44) that were not used as training data for the M-distance
model. This analysis showed that the distribution of nests within each habitat
category fit the expected distributions (v = 3, X2 = 1.02, P =
0.80; Table 1). This suggests that the simple rules did a good job at defining
potential
habitat, and that M-distance was effective at predicting the area within
potential habitat where a given percentage of nests would be expected
to occur.
Table 1: Distribution of Mexican Spotted Owl nests
(n = 44) in the test dataset in relation to categories generated using
Mahalanobis distances and the nests in the training datasets.
M Distance
Range
|
Category
|
Expected % of Nests in Range
|
Actual Number and Percent of Nests in Range
|
1.4 - 5.1
|
4
|
50%
|
25 (56.8%)
|
5.1 - 7.3
|
3
|
25%
|
11 (25.0%)
|
7.3 - 10.6
|
2
|
15%
|
4 (9.1%)
|
10.6 - 14.9
|
1
|
10%
|
4 (9.1%)
|
Sources of errors
The potential habitat model developed above contained
about 85% of the known Mexican Spotted Owl nest sites. There are several
potential reasons that some sites may fall outside of predicted habitat.
First, there is some uncertainty in the basal area and vegetation type
layers used in the modeling. Thus some areas that are suitable habitat
may not be identified as such. Second, many of the owl nest sites had
positioning errors of + 100m, so the location of some nests on the
landscape may be inaccurate. Third, conditions may have changed at
some nest sites since they were last active. All of these may lead
to errors.
It should also be noted that certain assumptions must be made when
using the M-distance statistic as a measure of habitat “quality” or “preference”.
The most important of these assumptions are that the distribution of habitat
characteristics at known nest sites is representative of the actual distribution
of nest sites, and that the mean habitat characteristics at nest sites represent “optimal” or “preferred” habitat
characteristics. We do not currently have the data to determine whether M-distance
values can be linked to survivorship or reproductive success, and thus we do
not know how good an estimator of habitat quality is provided by the M-distance
statistic.
Recommendations
We recommend that this layer be used at a minimum
resolution of 90m (0.8 ha or 2 acres) for purposes of analysis
and display. However, ForestERA data layers were not designed for analyses
at the level of individual pixels, and uncertainty in the data
will
generally decline over greater spatial extents. Therefore, we recommend
using larger analysis units, with groupings of at least 50 cells
(40 ha or 100 acres). Finally, we reiterate that ForestERA data layers
were developed for the purpose of regional landscape-level planning,
and we suggest that the analyses be applied over spatial extents
of
tens to hundreds of thousands of acres. We recognize, however,
that this layer may be useful for analyses over smaller spatial extents
depending on the type and purpose of those analyses.
Literature Cited
Clark, J. D., J. E. Dunn, and K. G. Smith. 1983. A multivariate
model of female black bear habitat use for a Geographic Information
System. Journal of Wildlife Management 57: 519-526.
Corsi, F., E. Dupre, and L. Boitani. 1999. A large-scale model of
wolf distribution in Italy for conservation planning. Conservation
Biology 13: 150-159.
Farber, O. and R. Kadmon. 2003. Assessment of alternative approaches
for bioclimatic modeling with special emphasis on the M-distance. Ecological
Modelling 160: 115-130.
Ganey, J. L. and R. P. Balda. 1994. Habitat selection by Mexican Spotted
Owls in northern Arizona. Auk 17: 162-169.
Grubb, T. L., J. L. Ganey, and S. R. Masek. 1997. Canopy closure around
nest sites of Mexican Spotted Owls in northcentral Arizona. Journal
of Wildlife Management 61: 336-342.
Knick, S. T. and D. L. Dyer. 1997. Distribution of Black-tailed Jackrabbit
habitat determined by GIS in southwestern Idaho. Journal of Wildlife
Management 61: 75-85.
May, C. A. and R. J. Gutierrez. Habitat associations of Mexican Spotted
Owl nest and roost sites in central Arizona. Wilson Bulletin 114: 457-466.
McComb, W. C., M. T. McGrath, T. A. Spies, and D. Vesely. 2002. Models
for mapping potential habitat at landscape scales: an example using
Northern Spotted Owls. Forest Science 48: 203-216.
Prather, J. W., J. L. Ganey, P. Beier, W. M. Block, M. Ingraldi, J.
Jenness, Y. Xu, H. M. Hampton, and T. D. Sisk. (in review). Simple
models successfully map nesting habitat for two raptors in southwestern
coniferous forests. Conservation Biology.
Seamans, M. E. and R. J. Gutiérrez. 1995. Breeding habitat
characteristics of the Mexican Spotted Owl in the Tularosa Mountains,
New Mexico. Condor 97: 944-952.
U. S. Fish and Wildlife Service (USFWS). 1995. "Recovery plan for
the Mexican Spotted Owl (Strix occidentalis lucida)." U. S.
Fish and Wildlife
Service, Albuquerque, New Mexico.
Young, K. E., R. Valdez, P. J. Swank, and W. R. Gould. 1998. Density
and roost site characteristics of Spotted Owls in the Sierra Madre
Occidental, Chihuahua, Mexico. Condor 100: 732-736.
Zar, J. H. 1999. Biostatistical Analysis, 4th Edition. Prentice-Hall
Inc. Englewood Cliffs, New Jersey.
Zwank, P. J., K. W. Kroel, D. M. Levin, G. M. Southward, and R. C.
Romme. 1993. Habitat characteristics of Mexican Spotted Owls in southern
New Mexico. Journal of Field Ornithology 65: 324-334.
Last updated
March 4, 2005
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