Ethology Ecology & Evolution 12: 405-417, 2000

Factors affecting habitat use by mule deer

(Odocoileus hemionus) in the central part of the
Chihuahuan Desert, Mexico: an assessment with
univariate and multivariate methods

G. SÁNCHEZ-ROJAS 1,2 and S. GALLINA 2

1 Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Hidalgo,

Apartado Postal 69 Plaza Juárez, Pachuca, Hidalgo, México 42001

(E-mail: gsanchez@uaeh.reduaeh.mx)
2 Instituto de Ecología, A.C., Departamento Ecología y Comportamiento Animal,

Apartado Postal 63, Xalapa, Veracruz, México 91000

Received 14 September 1999, accepted 30 June 2000

Habitat use by mule deer was studied in the Mapimí Biosphere Reserve in
the Chihuahuan Desert, Mexico. The deer population in this area is distributed
among isolated hills and sierra. Because of their similar soil and vegetation
characteristics, these hills are a homogeneous element of the landscape. This
study was carried out within this element, in order to understand the habitat
variables that are important for deer in the selection of habitat. Univariate as
well as multivariate analytical methods were used in a complementary fashion to
identify habitat variables associated with habitat selection during wet and dry
seasons. The univariate method detected a relationship between habitat use and
variation in slope (measurements of unevenness in terrain), in both seasons, it
also detected a relationship between habitat use and the distance to water in the
dry season. The multivariate method detected higher use of sites with greater
unevenness in the terrain, higher shrub diversity, greater visibility and that were
closer to water sources. Unevenness in terrain seems to be the most important
factor influencing habitat selection by mule deer. This factor is limited to isolat-
ed hills and sierra in this part of the Chihuahuan Desert, so it is a spatially well
defined resource. Such areas should be considered as a high-priority for the con-
servation of this species.

KEY WORDS: Odocoileus hemionus, population, habitat use, conservation, bios-

phere reserve, ordination, multiple regression, landscape element.

Introduction . . . . . . . . . . . . . . . . . 406
Methods . . . . . . . . . . . . . . . . . . 407
Results . . . . . . . . . . . . . . . . . . 409
Discussion . . . . . . . . . . . . . . . . . 412
Acknowledgments . . . . . . . . . . . . . . . 415
406 G. Sánchez-Rojas and S. Gallina

References . . . . . . . . . . . . . . . . . 415
Appendix . . . . . . . . . . . . . . . . . 416

INTRODUCTION

The study of habitat selection by populations of wildlife continues to play an

important role in the effective management of their populations (OTIS 1997). The
term “selection” is not synonymous with the term “preference”. In this study, we
define “selection” as the process by which an animal chooses a resource and “pref-
erence” as the probability that a resource might be selected if offered under the
same conditions as other resources (JOHNSON 1980). Thus, habitat use is the result
of the selection process.
Habitat selection is generally studied via two approaches. The first approach
considers habitats as discrete categories (e.g., open fields, forest, rocky areas, types
of vegetation) in which the availability of the resource is taken into account with
respect to the use that an animal makes of it (in this case, the availability is usually
expressed as function of area). The second approach is to consider the attributes of
the habitat as a continuum (e.g., shrub density, percent cover, distances to water,
slope, etc.). Correlation between habitat values distributed along this continuum
and the intensity of use by an animal are measured (MANLEY et al. 1993).
When selection is evaluated using a continuum of habitat attributes, numer-
ous methods exist to analyze this information. Two of the most often employed,
are: (1) multiple regression analysis, in which a correlation might be established
between the dependent variable (usually a measure of wildlife populations such as
direct observations, tracks, fecal pellet group or location with radio-tracking) and
single habitat factor or combination of factors. (2) Multivariate methods, which
consider all variables at once and thus, permit a global description of the data. We
considered that multiple regression analysis is a univariate method because this
technique involves trying to account for variation in one dependent variable; and
we considered that multivariate analysis is when several related random variables
are considered simultaneously, each one being considered equally important at the
beginning of the analysis (MANLY 1997).
Although many analyses only involve one of these two methods, it has been
shown that both methods are complementary when used together. Accordingly, their
joint use might help us to better understand which factors are most likely to the
affecting the process of habitat selection by animals (BEN-SHAHAR & SKINNER 1988).
The distribution and abundance of deer populations, as well as those of other
ungulates, are affected by the availability of food, cover, and freestanding water in
the habitat because these resources provide the basic physiological and biological
requirements of the deer (DASMANN 1971). However, we currently recognize that in
order to understand the process by which ungulates select a habitat, it is necessary
to interpret them as a function of how these same factors (food, cover and water)
contribute to an increase in feeding efficiency and/or decrease in the risk of preda-
tion (MOLVAR & BOWYER 1994, BLEICH et al. 1997, BOWYER et al. 1998). According to
this conceptual framework, individuals would resolve the conflict between the
necessity to feed efficiently and the imperative necessity to avoid predation by
selecting habitats that minimize the ratio between predation risk and foraging effi-
ciency (PULLMAN 1989). The objective of this research was to determine the habitat
Habitat use by mule deer in Chihuahuan Desert 407

characteristics associated with habitat selection in two different seasons by a popu-

lation of mule deer, using uni- and multivariate methods.

METHODS

Study area

The Mapimí Biosphere Reserve (MBR) is located in the northeastern part of the state of
Durango, Mexico, bordering the states of Chihuahua and Coahuila. The reserve is included
within the Man and Biosphere Program (MAB-UNESCO) and was established with the goal of
preserving a representative portion of the typical ecosystem in the southern portion of the
Chihuahuan Desert. The MBR polygonal boundaries are located between 26°48’ and 26°31’N
and 104°03’ and 103°32’E (Fig. 1).
The MBR is located in the Mapimí Bolson, which is a depression or closed basin into
which rivers and streams flow during the rainy season. The highest point in the area is San
Ignacio Hill 1480 m a.s.l., and the average altitude of the basin floor in the reserve is approxi-
mately 150 m. Broad plains and small mountain ranges characterize the region. In the MBR
(1720 km2) seven landscape elements were identified: bajadas and hills of igneous and sedi-
mentary origin (37% of the area); bajadas and sierra of calcareous origin (17%); southern
playa (18%); northern playa (6%), dunes (13%); eolian-fluvial transition (8%); and basaltic
lava flows (1%) (MONTAÑA & BREIMER 1988).

Fig. 1. — Location of the Mapimí Biosphere Reserve, including the two samples sites.
408 G. Sánchez-Rojas and S. Gallina

Average annual precipitation was 271 mm (based upon 25 years of data). The wet sea-
son occured from July to December, and the dry season from January to June. Average annu-
al temperature was 20 °C with a seasonal variation of 16 ºC and daily variation of 20 ºC
(MONTAÑA 1990). Based upon information provided by the climatological station at MBR (L.
HERNÁNDEZ unpublished data), precipitation was significantly different (t = 2.209; df = 1; P =
0.048) between the two cycles of the dry and wet seasons during the time of this study, while
temperature did not vary significantly (t = 0.211; df = 1; P = 0.835).
In a preliminary survey carried out within permanent plots for 6 months to determine
the distribution of mule deer within the landscape elements in the reserve, we observed indi-
viduals or their signs (tracks and pellet groups) only in the bajadas and hills of igneous and
sedimentary origin and bajada and sierra of calcareous origin (SÁNCHEZ-ROJAS & GALLINA
2000). Thus, we consider that currently only these landscape elements provide mule deer with
a suitable habitat, these elements being embedded within a matrix of flatlands and plains.
Due to the need of establishing management strategies for this species within the MBR,
it is important to know the factors within the element of bajadas and hills of igneous and
sedimentary origin that influence habitat use by deer. The element of bajadas and hills of
igneous and sedimentary origin are located in the central and west-central part of MBR. The
vegetation is dominated by Larrea tridentata and Fouquieria splendens (or less frequently, L.
tridentata and Yucca rigida). In some areas, succulent species (Opuntia microdasys, O.
rastrera, Agave asperrima, and A. lechuguilla) co-dominate with L. tridentata. The bajadas and
hills are shallow and/or rocky (MONTAÑA & BREIMER 1988).

Data collection

To include the greatest variation in the habitat attributes of the bajadas and hills of
igneous and sedimentary origin, two sites were utilized which are known to belong to this
landscape element (SÁNCHEZ-ROJAS & GALLINA 2000), one in San Ignacio Hill and the other in
Coronas Hill (Fig. 1).
A total of 14 permanent transects were established, 8 in San Ignacio Hill and 6 in Coro-
nas Hill. The starting point and orientation of each transect were randomly chosen, each
transect measured 780 m length, and within each transect 40 circular sampling plots were
uniformly distributed, with a radius of 3 m. Within each plot, all fecal pellet groups were
recorded for each sampling event and the wet or dry season was noted. In this study, we con-
sidered the sampling events of July and September of 1996, and September and December of
1997, as representing the wet season. The sampling events of March and June of 1997 and
1998 represented the dry season.
In order to characterize each transect we considered 12 habitat variables: (1) shrub
density; (2) shrub height; (3) shrub crown; (4) visibility index; (5) total cover of species con-
sumed by the mule deer; (6) total cover of species not consumed; (7) average distance to
water; (8) minimal distance to water; (9) variance of slope; (10) slope; (11) shrub diversity
index using the cover value; (12) shrub diversity index using the frequency. The methods used
to obtain the measures appear in Appendix.

Statistical analyses

A three-way analysis of variance was performed in which the total number of pellet
groups collected was compared among transects, seasons and the two sampling cycles of the
dry and wet seasons; the pellet count data was square root transformed (ZAR 1996).
Both sites were pooled as a unit, to increase the variation in the condition of the 12
habitat characteristics used. Thus, we believed that the conditions encountered toward the
interior of the landscape element could be better generalized.
With the data matrix of pellet counts as the dependent variable and the value of the 12
habitat variables as independent variables, a step-wise forward multiple regression was per-
Habitat use by mule deer in Chihuahuan Desert 409

formed in order to determine which of the 12 habitat variables were significantly associated
with the number of fecal pellet groups to avoid multicollinearity (ZAR 1996). When the vari-
ance was not homogeneous or residuals were not normally distributed, the values of the vari-
ables were transformed.
An ordination procedure of the transect was performed, using these 12 habitat charac-
teristics. This ordination relied upon a principal component analysis (PCA) procedure which
permits a complex set of original variables to be reduced into a smaller set of derived inde-
pendent factors (components). The analysis used standardized variables since the original
variables were expressed in different units (RANDERSON 1996). Pearson correlation coefficients
were used to establish which variables were significantly associated with each component
(MORRISON et al. 1992).
Once the value for each one of the transects in the components was obtained, this was
used as an independent variable in a linear regression against the number of pellet groups. In
this way, it could be established if there was a pattern between the biological interpretation
assigned to each component and the use that the population made of the different condition
of the same landscape element.

RESULTS

The analysis of variance of fecal pellet groups showed that there are signifi-
cant differences among transects (F = 9.806; df = 13; P < 0.001) and between the
wet and dry season (F = 8.931; df = 1; P = 0.010), but there were no significant dif-
ferences between the 2 years sampled. For this reason, we pooled the information
over both years considering the number of pellet groups as dependent variables
with values in each transect between each seasons.
Table 1 contains estimates for each one of the 12 habitat characteristics, as
well as the principal component values that correspond to each transect in both
sites, and the pellet count data by season. We made a logarithmic transformation
with the value of variance of slope and the average distance to water, to perform
the regression analysis.
The multiple regression of pellet groups against the different variables in the
wet season shows a significant association only with the variance in slope (F =
25.488; df = 1; P < 0.001), having a value of r2 = 0.680 (Table 2). In the dry season
the significantly associated variables were the variance in slope (F = 31.602; df = 1;
P < 0.001) in step one of the regression and the average distance to water (F =
4.632; df = 2; P = 0.05 in step two; the combination of this two variables having a
value of r2 = 0. 813 (Table 2).
The PCA analysis assigns 66% of the accumulated variance to the first two
principal components (Table 3). The results of the correlation analysis, between the
values of the habitat variables and the component values for each transect, allowed
us to objectively select which combination of these habitat variables were more
important in each component. Importance was judged according to significance
level (yes or no) of the correlation (Table 3).
Upon completing the regression between the number of pellet groups and the
values of the transects from the ordination of the two components, we found that
only component I had a relationship with habitat use. Component I was positively
correlated with shrub density, visibility index, slope, variance in slope and plant
diversity based on cover and frequency; and was negatively correlated with shrub
crown size, shrub height, the quantity of non-consumed plants, the average dis-
tance to water, and the minimum distance to water (Table 3).
 Table 1.
410

Values for habitat characteristics, the values for each transect in the first two components of the ordination analysis, and the number of fecal pellet
group in each sampling unit in the bajadas and hill of igneous and sedimentary origin within the Mapimí Biosphere Reserve. The first eight tran-
sects are from San Ignacio Hill, the last six are taken from Coronas Hills.

Transects

Variable San Ignacio Hill Coronas Hill

1 2 3 4 5 6 7 8 1 2 3 4 5 6

Shrub density /100 m2 33 21 42 30 24 42 25 8 20 20 18 15 28 26

Height (m) 0.71 0.97 0.76 0.89 0.85 0.88 0.84 0.79 1.06 0.93 1.06 1.00 0.97 0.93

Crown (m2) 4.72 5.81 5.09 5.94 5.47 5.51 5.69 5.8 7.61 6.25 6.9 6.98 5.80 5.70

Visibility index 1.76 2.17 2.33 1.72 1.79 2.08 1.82 1.82 1.47 1.48 1.89 1.59 1.79 1.47

Total of species consumed (m3) 588 637 637 490 361 394 544 459 624 566 565 603 376 502

Total of species not consumed (m3) 168 294 179 461 515 493 367 469 593 433 540 325 741 411

Average distance to water (km) 3.68 3.28 3.11 3.02 3.16 3.06 2.99 3.45 5.48 5.44 5.68 5.54 5.69 5.46

Minimal distance to water (km) 2.57 1.95 1.53 0.73 0.44 0.59 1.14 2.64 4.10 2.67 2.12 2.49 2.93 2.95

Variance of slope 10.97 0.28 57.4 4.02 15.78 29.40 29.47 0.12 1.08 1.79 11.11 0.13 1.73 4.05

Slope (degrees) 0.01 0.24 5.40 0.31 0.35 3.13 4.51 0.08 0.69 0.40 2.88 0.14 1.72 4.05

Shannon index (cover) 2.22 2.20 2.40 1.70 1.60 1.89 2.05 1.75 1.77 2.02 1.90 1.51 2.09 1.94

Shannon index (frequency) 2.27 2.18 2.41 1.83 1.76 2.08 2.12 1.78 2.03 2.14 2.07 1.87 2.26 2.29

Value of Component I 0.60 0.17 1.47 – 0.10 – 0.04 0.60 0.52 – 0.33 – 0.97 – 0.41 – 0.43 – 0.80 – 0.27 0.01

Value of Component II 0.17 0.17 0.46 – 0.61 – 0.93 – 0.44 – 0.04 – 0.49 0.48 0.29 0.30 0.04 0.16 0.43

Pellet groups in dry season 21 5 34 31 25 26 29 0 10 13 6 1 11 27

Pellet groups in wet season 19 19 29 28 38 41 48 1 19 24 22 0 26 29

G. Sánchez-Rojas and S. Gallina
Habitat use by mule deer in Chihuahuan Desert 411

Table 2.
Summary table of the results of a step-wise forward multiple regression, in wet and dry season, by
the pellets group counts and 12 habitat variables (for more details see text), SE is the standard
error.

Season Group Partial SE of SE F-Remove P R R2

regression coefficient
coefficients (β)

Wet Constant 17.458 2.520

Variance of slope 5.268 0.825 1.043 25.488 < 0.001 0.825 0.680

Dry Constant 111.905 47.134

Variance of slope 4.336 0.769 0.771 31.602 < 0.001 0.857 0.734
Average distance
to water – 12.111 – 0.294 5.632 4.624 0.05 0.901 0.813

Fig. 2. — Regression between the number of pellet groups and the value of Component I for
the ordination of 14 transect located on the slopes of igneous origin in the Mapimí Biosphere
Reserve. (A) Regression line for the rainy season; (B) Regression line for the dry season. In
both cases the regression equation is included in which “y” is equal to the number of pellet
group and “x” is equal to the value on Component I.
412 G. Sánchez-Rojas and S. Gallina

Table 3.
Results of the ordination of the variables in bajadas and hills of igneous and sedimentary origin,
using a PCA. The Pearson correlation coefficient was used to estimated the association between the
values of each variable along the component axes and the original value of variable (* = P < 0.05,
** = P < 0.001, – = negative correlation, + = positive correlation).

Component I Component II

Variation explained (%) 44.43 21.13

Cumulative variation (%) 44.43 65.74

Variable Correlation with original variables

Shrub density /100 m2 + ** NS

Height (m) – ** NS
Crown (m2) – ** NS
Visibility index +* NS
Total of species consumed (m3) NS +*
Total of species not consumed (m3) –* NS
Average distance to water (km) –* +*
Minimal distance to water (km) –* +*
Variance of slope + ** NS
Slope (degrees) +* NS
Shannon index (cover) +* +*
Shannon index (frequency) +* +*

The relationship with the transect value for Component I against the number
of pellet groups has a marginally significant slope in the rainy season (F = 4.83; df
= 1; P = 0.048), with a low coefficient of determination r2 = 0.287 (Fig. 2A). In the
dry season the relationship has a significant slope (F = 11.97; df = 1; P = 0.005),
having a coefficient of determination value of r2 = 0.499 (Fig. 2B).

DISCUSSION

We employed fecal pellet group counts to determine the activity patterns of

mule deer, and therefore the use of habitat by this ungulate. It has been previously
questioned whether the distribution of pellet groups accurately reflects habitat use
(COLLINS 1981, COLLINS & URNESS 1984, LEOPOLD et al. 1984). However, it has been
argued that in those studies in which the use of pellet groups has been compared,
at the same time, with some other technique (observation, tracks and/or radio
tracking) in order to estimate the relative use of habitats, the results did not differ
significantly between methods (CAIRNS & TELFER 1980, LEOPOLD et al. 1984, LOFT &
KIE 1988). The method of pellet groups counts could have certain problems of
interpretation, specifically, an increase in the number of pellet groups does not nec-
essarily imply an increase in the density of animals. Rather, it indicates an
increased presence or increase in activity time (WALLMO 1969, KIE 1984).
Habitat use by mule deer in Chihuahuan Desert 413

The results of the two analyses used ih the present study have different
assumptions and calculation procedures. While the multiple regression analysis cal-
culates the correlation between various parameters and assigns probability levels to
these, the ordination analysis provides us with a non-parametric description of the
relationship between the variables and is an indicator of importance more than a
measure of significance (BEN-SHAHAR & SKINNER 1988)
The result of the regression analysis shows that there is a significant relation-
ship (P = < 0.001) in the wet season between the use of the habitat and the uneven-
ness of the terrain, while in dry season habitat use had significant relationships (P
= < 0.001) with the unevenness of the terrain and the average distance to the water.
Considering this, we suppose that only these variables can explain habitat use by
mule deer. However, species often respond to an environmental variable or gradient
in a non-linear fashion (JOHNSON 1981).
The PCA analysis, produces “new” variables that are the linear combination of
the original variables. The variable represented by the Component I is the only one
that shows a relationship with habitat use by mule deer. In this case, we assume
that the regression between this variable and the use of the habitat provides us
with an objective method of deciding if there was a pattern. If there is a significant
slope, then the pattern exists, and the sign of this slope indicates the pattern’s
direction. In this case, we found that the slope is significant, and its sign is positive
in both seasons.
Our biological interpretation of Component I is that of a continuous habitat,
where the greater values in Component I correspond to transects with a greater
unevenness in terrain, higher plant density, higher diversity, greater visibility, small-
er size in the plants and a shorter distance to the sources of water. The other
extreme of this continuum corresponds to transects where a lower unevenness in
terrain, lower plant density, lower diversity of plants, the size of the plants is high-
er, and the sources of water were more distant.
Our viewpoint is that if we accept only the results of the multiple regression,
we could simplify the variables that are associated to the process of habitat selec-
tion by mule deer. Otherwise, if we also use the results of the multivariate method,
we would additionally find that there are other important variables to understand
the selection of the habitat, which might not be considered because they are not
statistically significant, although these variables may have an important biological
meaning. The exploration of this biological meaning could be possible using avail-
able biological data on the mule deer.
GEIST (1998) provide a meticulous description of antipredator strategies by
mule deer. He describes six different strategies: (1) stotting (particular gait of mule
deer, that are long high and fast bounds, which is more efficient than a speedy gal-
lop, when travelling up hill); (2) random daily movement; (3) hiding (but in differ-
ent form from white-tailed deer, get-aways); (4) detecting danger at long distances
and vacating the areas; (5) attacking predators and (6) forming herds (GEIST 1998,
see references in his paper).
The results of our study agree with the strategies described by GEIST (1998).
We found that mule deer select areas with uneven terrain, because the high trajec-
tory of the mule deer’s bound allows them to ascend slopes easily. Ascending
straight up in an uneven terrain would be a formidable obstacle to any predator
that gallops (GEIST 1981, 1998). This selection by uneven terrain has been observed
in other populations of mule deer (RILEY & DOOD 1984, CARSON & PEEK 1987, GALLI-
NA et al. 1991, FOX & KRAUSMAN 1994).
414 G. Sánchez-Rojas and S. Gallina

The multivariate analysis showed that the transect with more unevenness of
terrain additionally had more visibility. This combination of factors is perhaps
important in habitat selection by mule deer, because deer prefer areas with more
visibility, to detect an approaching predator at long distances, so that they can
avoid the predator by moving off (GEIST 1998).
The results also showed that the average distance to a water source is an
important factor to consider, principally in the dry season, since deer generally
require a source of the standing water in order to survive periods of water stress
(HERVERT & KRAUSMAN 1986). Thus, the distance to water appears to affect negative-
ly habitat use by mule deer. This dependency on water has also been detected in
populations that live in desert environments (BOWYER 1984, HERVERT & KRAUSMAN
1986, ORDWAY & KRAUSMAN 1986).
The variable associated with food availability was not informative using either
of the two methods. In fact, in this study, the availability of plants consumed by
deer does not differ significant from the bajadas and hills of igneous origin. This
could be because mule deer respond more to the quality, rather than the quantity
of food (WECKERLY 1994). However, the multivariate method showed that diversity
and density of the woody plants and the succulent strata also seem to be important
given that there was a greater use of transects that offered a higher diversity and
density of vegetation, together with areas with more unevenness in the terrain.
Mule deer consume a great variety of plants in small amounts (KRAUSMAN et al.
1997). In addition, an association between greater use and increased shrub density
has been documented in other populations of mule deer in the Chihuahuan Desert
(LEOPOLD & KRAUSMAN 1991).
Perhaps the importance of both plant diversity and density can be explained
as a reflection of the need for greater variety of plants in order to increase foraging
efficiency and thereby balance the nutritional requirements of the deer. Because of
this, we think that detecting a greater presence of mule deer in areas with higher
diversity and greater density of shrubs could serve as indicator of the quality of the
site, at least in terms of food, since the deer would have a greater opportunity to
select better food among a greater number of plant species.
In order to understand the process by which mule deer select habitat, we
need to interpret the factors that contribute to an increase in feeding efficiency
and/or decrease in the risk of predation (MOLVAR & BOWYER 1994, BLEICH et al.
1997, BOWYER et al. 1998). In MBR mule deer seem to select the places where the
risk of predation is minimized. These places also coincided in being closest to
water and in having a combination of more diversity and density of plants. These
factors could favor the feeding efficiency of the mule deer. Thus there is no conflict
between feeding efficiency and the risk of predation, because the safest places
could also increase the feeding efficiency of mule deer. The deer only select the best
places and do not need to make compromisses between these factors.
These zones of greater activity by mule deer are located among the isolated
hills of the MBR and they are generally found at intermediate altitudes. Unfortu-
nately the local people have the common practice of using these zones as feeding
ground for cattle at the end of the dry season, since they offer little available forage
(principally shrubs and succulents) (BARRAL 1988). The perturbation of these zones
could have consequences for the future of the mule deer population, since it is a
spatially limited resource.
Habitat use by mule deer in Chihuahuan Desert 415

ACKNOWLEDGMENTS

The authors thank L. Hernández, V. Sánchez-Cordero, S. Mandujano, C. Moreno, R.

Macías, R. Guevara J. Heffelfinger, J. Loundré, A. Meriggi and one anonymous reviewer for
their comments, suggestions and ideas on the first draft of the manuscript. Also thanks to the
Herrera family for their help in the Reserve. We thank Dan Bennack for the translation into
English. The study was supported by CONACyT (The National Council of Science and Tech-
nology) as part of project 225660-5-2480PB to S. Gallina. In addition, CONACyT provided a
study scholarship and Fondo Mexicano para la Conservación de la Naturaleza awarded a
research grant to G. Sánchez-Rojas.

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APPENDIX

Habitat measurement

In order to characterize the vegetation along each transect, we used the “point-centered
quadrant” method (MUELLER-DOMBOIS & ELLENBERG 1974), considering only the woody and
succulent plants. The same 40 points where pellet groups were collected were also used to
sample the local vegetation. A total of 160 plants were measured along each transect. From
Habitat use by mule deer in Chihuahuan Desert 417

each plant, the following measurements were taken: distance to the central point of the quad-
rants, height of the plant, and diameter of the crown. With this information, the density, area
of crown, and volume were calculated for every plant. Additionally with the values for dis-
tance, shrub density per 100 m2 was estimated since the total density (TD) was approximated
as TD = 100/d2 where d is the average distance of the plant to middle point obtained for each
transect (MUELLER-DOMBOIS & ELLENBERG 1974). Every one of the plants measured was identi-
fied to the species level. With the value of frequency and cover for each species, the Shannon
index was calculated as a measure of diversity. In order to separate the volumes of potential
mule deer forage plants from those plants that are not eaten, we used the available informa-
tion on mule deer diets in the Chihuahuan Desert (GUTH 1987, KRAUSMAN et al. 1997). We
report the total volume of each group of plants.
Hiding cover was estimated according to the method suggested by GRIFFITH & YOUTIE
(1988), using a 2 m × 5 cm bar alternately painted with black and white stripes, each measur-
ing 10 cm in length. The bar was placed at the distance of 15 m perpendicularly from the
center point of the sample plot on either side of the transect. From the center point of each
plot, the percent area of stripes that were visible to an observer on one knee was estimated.
This height of observation was judged to be similar to the visual height of the deer. With this
percentage hiding cover, an index of visibility was estimated. This index was calculated as the
inverse of hidden cover so that a great index value correspond to less hidden cover (BOWYER
et al. 1998).
The coordinates of the transect locations, and water sources found around the study
area, were determined with a Garmin 45® geographic geopositioning device. With this infor-
mation, the distance from water sources to the mid-point of each transect was calculated.
Two values were obtained: the average distance from the transect to all sources of water in
each locality, and the minimum distance.
For each transect we used a clinometer (Suunto®) to estimate the degree of slope along
the 20 m intervals separating each sampling plot. These data were used to calculate the aver-
age slope and its variance for each transect. Both values were taken to represent measure-
ments of the unevenness of the terrain.