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Xeroriparian Systems Used by Desert Mule Deer
in Texas And Arizona 1
Paul R. Krausman 2 , Kurt R. Rautenstrauch 3 and Bruce D. Leopold 4
Abstract.--We examined desert mule deer (Odocoileus
hemionus crooki) occurrance in xeroriparian systems in
Arizona and Texas.
Most deer in Arizona were located in
washes. Most deer in Texas were located between washes.
Xeroriparian areas are important habitat components for
desert mule deer when they provide forage, thermal cover and
travel lanes.
INTRODUCTION
STUDY AREAS
Desert mule deer inhabit the Sonoran and
Chihuahuan Deserts of North America.
Their range
extends from southwest Texas to western Arizona
and south into central Mexico (Wallmo 1981).
Desert mule deer use of xeroriparian systems
was evaluated on the northeastern edge of their
range in Big Bend National Park (BBNP), southwest
Texas; in the westcentral part of their range in
the Belmont Mountains, central Arizona; and on the
northwestern edge of their range in King Valley,
southwest Arizona.
Desert mule deer are a popular and important
game animal, but have received limited attention
by
the scientific community.
Clark
(1953)
examined desert mule deer behavior and movement
patterns, Truett (1972) studied their general
ecology,
Krausman (1978) and Leopold
(1984)
evaluated their forage preferences, and Krausman
(1984) and Rautenstrauch and Krausman (unpublished
data) have studied desert mule deer home range
size and movements.
Descriptions of desert mule
deer habitat are general (Phillips 1974, Anthony
and Smith 1977, Dickinson and Garner 1979, Koerth
1981, Leopold and Krausman (1983) and there is
little published information on desert mule deer
habitat use.
BBNP, Brewster Co., is representative of the
rugged Chihuahuan Desert and is included in the
Chisos biotic district (Dice 1943).
Elevations
extend from 573 m along the Rio Grande to 2384 m
at Mt. Emory in the Chisos Mountains.
BBNP is characterized by hot summers, mild
winters and low rainfall.
Temperatures exceed 38
C in the desert regions in summer and rarely
freeze in winter.
Precipitation occurs from May
through October, ranging from 28-41 cm.
Leopold and Krausman (1983) identified 10
vegetative associations in BBNP. The associations
were differentiated into three categories based on
dominant
plant
cover:
creosotebush
(Larrea
tridentata) dominated, non-creosotebush dominated,
and associations not dominated by shrubs.
We began studying desert mule deer in Texas
in 1972 (Krausman and Ables 1981, Leopold 1984),
and
in
Arizona
in
1979
(Krausman
1984,
Rautenstrauch and Krausman, unpublished data).
During these studies it became apparent that
xeroriparian
washes
and
their
associated
vegetation were an important component of desert
mule deer habitat.
Our objective in this study
was
to
document desert mule deer
use
of
xeroriparian systems (Johnson et ale 1981) across
the northern boundary of their range and to
describe the vegetation of washes used by deer.
The Belmont Mountains, Maricopa Co., are 80
km west of Phoenix, cover 360 km2 , and are
representative
of the upper Sonoran
Desert.
Elevations range from 426 m to 914 m.
The average annual precipitation is 20 cm.
Most rain falls from January through
March.
Temperatures above 45 C in summer are common.
1Paper presented at the first North American
Riparian Conference.
[University of Arizona,
Tucson. April 16-18, 1985].
2Paul R. Krausman is Associate Professor of
Wildlife, University of Arizona, Tucson, Az.
3Kurt R. Rautenstrauch is Graduate Research
Assi~tant, University of Arizona, Tucson, Az.
Leopold is Wildlife
Research
Bruce D.
Assistant, University of Arizona, Tucson, Az.
Krausman
(1984) identified 9
vegetative
associations in the Belmont Mountains.
Most
associations are dominated by triangleleaf bursage
(Ambrosia
deltoidea),
brittlebush
(Encelia
farinosa) and creosotebush. The areas between the
mountains and foothills having major washes are
classified as the Triangleleaf Bursage-Transition
Association.
Vegetation in this association is
dominated by the same three plants, but the washes
144
contain the larger trees, ironwood
and paloverde (Cercidium sPP.).
(~
Arizona
tesota)
Belmont Mountains.--The density of perennial
vegetation
was measured in
50-100
0.004-ha
randomly located Circular plots in each of nine
associations identified.
In major washes that
bisected associations, line intercept transects
(Canfield
1941) were established to estimate
the
vegetational composition in
xeroriparian
components of the association.
Over 60% of the Belmont Mountain area is in
the Creosote Flats Association.
Dominant plants
include creosotebush and triangleleaf bursage.
The species composition of the major washes is
similar to the transition association.
King Valley, Yuma Co., is 45 to 60 km
northeast of Yuma and 110 kID southwest of the
Belmont Mountains.
Elevations range from 85 m at
the Gila River to 450 m at the base of the
surrounding
mountains.
The
average
annual
precipitation at the lower end of the valley is 12
cm.
King Valley.--Washes in King Valley were
divided into 4 classes depending on the number of
drainages
and the width of
the
associated
vegetation.
Simple washes have only one drainage
(a water fluve greater than 1 m wide).
Complex
washes (washes with more than 1 fluve) were
divjded into 3 classes based on the width of the
vegetation: less than 50 m wide (C1), between 50
and 150 m wide (C2) and greater than 150 m wide
(C3).
The slopes of the mountain ranges surrounding
King Valley are sparsely vegetated and dominated
creosotebush,
brittlebush,
white bursage
by
(Ambrosia
dumosa), and
ocotillo
(Fouquieria
splendens).
The canyon bottoms have xeroriparian
washes dominated by ironwood and paloverde.
Most plant life in King Valley is restricted
to the xeroriparian drainages.
The areas between
drainages are usually covered with wind eroded
desert pavement, and have no vegetation or very
sparse stands of creosotebush, brittlebush and
white bursage.
The dominant overstory species in the washes
are little-leaf paloverde (~ microphyllum), blue
paloverde (~ floridum) and ironwood.
The width
of the vegetation in the largest washes in King
Valley is over 300 m wide.
The percent cover of perennial vegetation in
washes
was measured using the line-intercept
method (Canfield 1941). Ten transects, spaced ten
m apart and running perpendicular to the flow of
water, were measured in each wash.
The density of perennial vegetation between
10 .0314 ha
washes was measured in six to
circular plots next to each wash measured.
The
plots were either 100 m away from the edge of the
wash vegetation or half-way between the measured
wash and the adjacent wash if the washes were less
than 200 m apart.
Contrasting Washes and Adjacent Habitat
Because deer were rarely located in the
mountains and foothills surrounding King Valley,
this study deals only with the
xeroriparian
Paloverde-Ironwood Association found in the bottom
of King Valley.
METHODS
Vegetation Sampling
Shannon-Weaver
Diversity
indices
were
computed to contrast plant species diversity in
washes and adjacent habitats in Texas and between
areas in Arizona.
Morisita coefficients
of
overlap (Morisita 1959) were computed to determine
degree
of similarity of perennial vegetation
within
washes
and
adjacent
vegetatiave
associations in Texas. The equivilence of percent
forage species occurring in the washes and the
adjacent vegetatiave associat!ons was determined
using the binomial test for proportions (Zar
1984:395-400).
Texas
Deer Occurrence in Washes
Twenty five to 50 point-quarter plots (Dix
1961) were sampled in each of ten vegetative
associations identified in BBNP (Leopold
and
of
Krausman
1983) to determine the density
vegetation between washes.
Texas
Deer use of washes was determined from 750
independent observations of deer in three classes:
initially observed in wash, within 30 m of a wash,
or
greater than 30 m from
a
wash.
All
observations were made from January 1980 through
1981.
Habitats
were
sampled for deer
in
proportion to their availability in the study
area.
A transect line was established along the
center of each sampled wash running parallel to
the flow of water.
The initial plot was randomly
determined and subsequent points were 15 m apart.
At each point, width of wash, and all perennial
plant species to the left and right of the point
were recorded.
We also noted which plants were
deer forage. Desirable deer forage plant species
was based on diets determined by fecal analysis
(Leopold 1984).
Arizona
Deer use of washes was determined from 1180
independent locations of 12 radio-collared deer (4
145
Table
1.--Summary and comparison of vegetative characteristics of plant associations and adjacent wash
systems in Big Bend National Park, Texas.
Plant association
Vegetative
association
Total Pltnt Deer
density
forage 2
Creosote bush dominated
3.01
Creo-Iech-grass
Creo-Iech-candel
0.74
0.65
Creo-Iech-Opuntia
Creo-tarbush
2.30
Creo Flats (Loc 1)4 0.64
Creo Flats (Loc 2)5 0.04
0.24
Creo-Iech
II. Non-creosotebush dominated
2.46
Vig-Iech-grass
Yucca-Sotol
4.34
Sotol-Iech-grass
4.05
III. Non-shrub dominated
Lech-grass (Loc 1)
1.72
Lech-grass (Loc 2)
1.04
vlash systems
Deer
forage 2
vlash
width 3
Diversity
Adjacent
habitat
wash
Coefficient
of overlap
I.
43.4*6
28.3
27.8
40.1*
5.9
0.0
33.6
29.3
41.3**7
31 .1
26.4
8.2
55.8**
55.3**
18.6
23.5
19.6
5.4
13.3
25.0
NA
2.25
2.31
1.59
2.69
1.55
1.09
1.95
2.81
2.38
2.58
2.25
1.80
2.37
1.59
0.09
0.40
0.27
0.50
0.66
0.30
0.32
44.3
46.8*
53.0
62.4**
23.7
60.2
5.5
9.7
6.3
2.58
2.74
1.87
2.40
2.70
2.87
0.76
0.37
0.25
68.6*
59.1
53.1
57.7
3.8
5.2
2.73
2.35
3.01
2.69
0.35
0.60
'expressed as stems/m2
2expressed as percentage
3expressed as average of all points sampled
4Creosotebush Flats of upper elevations
~Creosotebush Flats of lower elevations
*=deer forage in vegetation association significantly greater (alpha = 0.05) than in washes within
association
7**=deer forage in washes significantly greater (alpha = 0.05) than adjacent association.
males, 8 females) in the Belmont Mountains from
1980-1983, and 870 independent locations of 15
radio-collared deer in King Valley (4 males, 11
females)from 1982-1984.
Each collared animal was
located
weekly with a fixed
wing
aircraft
(Krausman et ale 1984).
For each location deer
were classified as being in a wash or not in a
wash. In King Valley the class of wash being used
was also recorded.
composition of adjacent habitats. This difference
was smallest in BBNP and greatest in King Valley.
Texas
Plant species within wash systems was not
similar to the perennial species composition of
adjacent habitats. Coefficients of overlap rarely
exceeded
0.60
(table
1)
which
represents
significant
biological overlap (Alcoze
and
Zimmerman 1973).
RESULTS
Plant species diversity was greater in washes
than in adjacent habitats for all but 4 plant
associations. As equal number of associations had
significantly greater forage percentages in washes
than in adjacent habitats (table 1).
The wash
systems in low density creosotebush dominated
associations were generally more diverse and had
greater deer forage percentages than the adjacaent
habitats.
Deer using plant associations with low
plant
densities
may therefore
find
higher
diversity and more forage in washes than in the
adjacent habitat.
Characteristics of Xeroriparian Systems
The average width of washes sampled in BBNP
ranged from 3.8 m to 25.0 m (table 1).
In
general, wash systems in the lower plant density
associations were wider than those with high plant
density associations.
Washes in the Belmont Mountains are similar
in size to those in BBNP.
The largest washes in
King Valley are wider than washes in the two other
study sites.
The average width of C washes is
3
284 m, and the largest washes measured were over
350 m wide.
Arizona
Belmont Mountains.--Plant species within wash
systems (tables 2, 3) was not similar to the
perennial
species
composition
of
adjacent
habitats. The plant species composition of washes
was more diverse than that of the surrounding
Contrasting Washes and Adjaceant Habitat
in
In all 3 study areas the species composition
wash systems was not similar to the species
146
Table
Table 2.--Vegetation in xeroriparian
systems
associated with the Triangleleaf bursageTransition Association, Belmont Mountains,
Arizona.
Plant species
% cover
Olneya tesota
Larrea trident at a
Cercidium microphyllum
Cercidium floridum
Prosopis juliflora
Lycium andersonii
Haplopappus larcifolius
Acacia greggii
Ambrosia ambrosioides
Ambrosia deltoidea
Condalia spathulata
Encelia farinosa
Hyptis emoryi
Krameria Er:ati.
Simmondsia chinensis
13.54
5.58
4.06
3.48
1.90
1.64
Total % cover
33.71
1.p
T
T
T
T
T
T
T
T
4.--Density (#/hectare) of perennial plants
in
the
Cresoste
Flats
(CF)
and
Triangleleaf
bursage-Transition
(TBT)
Associations, Belmont Mountains, and in the
Paloverde-Ironwood (PI) Association in King
Valley, Arizona.
Species
Belmont t-1ountains King Valley
TBT
CF
PI
Krameria spp.
Larrea tridentata
Carnegia gigantica
Opuntia spp.
Fouquiera splendens
Ambrosia dumosa
Ambrosia deltoides
Encelia farinosa
Other
12.5
280
12.5
500
12.5
2.5
1332.5
125
25
Total
Diversity
2302.5
= <1%
Plant species
417.5
35
25
13 .4
T
1232.5
141 .0
tesota
Cercidium microphyllum
Larrea tridentata
Cercidium floridum
Lycium andersonii
Ambrosia del to idea
Acacia greggii
Prosopis juliflora
Ambrosia ambrosioides
Acacia constricta
Haplopappus larcifolius
Condalia spathulata
Encelia farinosa
Fouguieria splendens
.fu:Q.lli ~
Krameria .ru:m
~
~spp.
Opuntia leptocaulis
Simmondsia chinensis
Sphaeralcea spp.
Total % cover
Diversity
1.31
1.01
1.00
Northern
washes
vegetation and provided a higher density of forage
and cover than adjacent areas.
King ~.--The
average
density
of
perennial vegetation in the habitat adjacent to
washes in King Valley was 1.4 plants/100 m2 (table
4).
These areas provide very little forage for
deer and have no shaded bedsites. Most preferred
forage species, such as ironwood, ratany, and blue
paloverde are uncommon or not found outside of the
washes.
Because the nonwash habitat, has no
overstory species and the common shrubs are small,
there are no shaded bedsites in these areas. Over
8% of the ground cover in washes is overstory
species that provided bedsites for deer (table 5).
Southern
washes
% cover
= <1%
T
4.2
T
10.4
1less than 1 plant/ha
cover)
systems
Table 3.--Vegetation
in
xeroriparian
Flats
associated
with
the
Creosote
Association, Belmont Mountains, Arizona.
1 T
T1
70
1.91
Diversity (H')
1 T
32.5
712.5
2.5
7.5
2.60
1.97
8.17
7.63
4.82
1.84
T
9.65
8.34
5.46
4.92
3.84
2.41
1.51
1.36
1.29
1.30
T
4
2·i
T
Deer Occurance in Washes
Texas
o
T
o
o
T
o
T
T
o
o
T
T
Arizona
T
Belmont Mountains.--Deer use of xeroriparian
systems was highest in summer (83.3%) followed by
fall (82.2%) and spring (70.5%) (table 6). During
the
winter deer use of washes
was
42.1%.
Overall, 842 of 1180 (71.4%) deer were located in
washes (table 6).
o
o
o
o
31.27
2.01
Of 750 deer observations only 40 (5.3%)
occurred within a wash, and 29 (3.9%) within 30 m
of a wash.
T
T
T
39.11
2.24
King ~.--Over 99% of the deer locations
in King Valley were in washes (table 7).
The six
locations that were not in washes were either in
cover)
147
Table 5.--Percent cover of vegetation in 4
classes in King Valley, Arizona.
~
Av.
Av.
Av.
Av.
Av.
n
..Q2.
wash
Table
7.--The number (and percent) of
deer
locations in 4 wash classes in King Valley,
Arizona •
n
Simple
width of wash
15.74 39.51 89.62 284.3
% cover of vegetation 30.40 29.36 23.76 24.77
II drainages
1.0
3.2
6.93 12.95
It of species
9.05 9.32 10.75
7.5
8.05
% of overstory cover 10.55 8.84 8.46
Hilaria rigid a
Atriplex polycarpa
0.05 1.02 0.08
0.28 0.94 0.11
0.19 0.35 0.41
0.30 0.10 0.03
0.96 0.25 0.13
3.54 1.88 1.42
1.42 2.55 3.28
5.10 3.96 3.32
10.48 11 .89 11 .05
0.18 0.31 0.23
3.60 2.85 2.56
3.12 1.18 0.65
2.57 2.94 1.67
0.86 1.28 1.09
Acacia~
Prosopis julif:lora
Krameria ~
Cercidium microphyllum
.Q... floridum
Olneya tesota
Larrea tr1dentata
Sphaeralcea SPp.
~ andersQnii
Ambrosia dumosa
Encelia farinQsa
Other species
Diversity (H')
2.10
2.12
1.93
0.56
0.79
0.52
0.25
0.24
5.64
0.81
9.97
2.52
3.47
0.16
0.55
1.38
n
C2
Other
£i
Females
47
(9.8)
184
62
172
(39.1)(36.6)(13.2)
5
( 1.1)
Males
32
(18.6)
44
28
67
(25.6)(39.0)(16.3)
1
(0.1)
In Arizona, most deer were located in washes.
III both Arizona study areas the plant species
diversity was twice as high in the xeroriparian
washes.
Food and cover was scarce outside of
these washes in King Valley and less abundant than
in washes in the Belmont Mountains.
Desert mule
deer in Arizona may be selecting xeroriparian
washes because they provide more food, cover, and
travel
lanes
than
the
surrounding
areas.
Xeroriparian systems are an important part of
desert mule deer habitat in xeric ranges.
2.03
LITERATURE CITED
agriculture
fields or disturbed
areas
agriculture at the south end of King Valley.
near
DISCUSSION
Deer in BBNP are not as dependent upon
xeroriparian systems as deer in Arizona.
In
Texas, deer forage is abundant in the habitats
adjacent to wash systems.
Although the plant
species
composition
of washes and
adjacent
habitats
are not similar,
both areas
have
relatively equal diversity,
except in
plant
associations with low plant densities.
Deer use
of
these areas was minimal «1.0
deer/km2)
compared to plant associations with higher plant
densities (>1.5/deer km 2 ) (Leopold 1984).
The
greater plant densities and diversities in the
interwash regions in BBNP allows deer to find
forage and cover in these areas instead of in
washes, as deer in Arizona must.
Alcoze, T. M., and E. G. Zimmerman. 1973. Food
habits and dietary overlap of two heteromyid
rodents from the mesquite plains of Texas.
J. Mammal. 54:900-908.
Anthony,
R.
G. ,
and N.
S. Smith.
1977.
Ecological relationships between mule deer
and
white-tailed
deer
in
southeastern
Arizona. Ecol. Monogr. 47:255-277.
Canfield, R. H.
1941. Application of the line
interception
method
in
sampling
range
vegetation. J. For. 39:388-394.
Clark, E. D. 1953. A study of the behavior and
movements of the Tucson Mountain mule deer.
M.S. Thesis, Univ. Arizona, Tucson.
111 p.
Dice, L. R. 1943. The biotic provinces of North
America.
Univ. Michigan Press, Ann Arbor.
78 p.
Dickinson, T. G., and G. W. Garner. 1979. Home
range use and movements of desert mule deer
in southwestern Texas. Proc. Ann. Conf. S.E.
Assoc. Fish and Wildl. Agencies 33:267-278.
Table 6.--Desert mule deer (4 males, 8 females) occurrence in washes in the Belmont Mountains, Arizona
from 1981-1982.
SEASONS
Jan-Mar
Habitat
Occurrences
Percent occurrences
in washes
Apr-Jun
Jul-Sep
Oct-Dec
Total
Wash
Other
Wash
Other
Wash
Other
Wash
Other
Wash
Other
82
113
322
135
304
61
134
29
842
338
42.1%
70.5%
83.3%
148
82.2%
71.4%
bighorn sheep.
Final Rept. 9-07-30-X069 to
USDI/B.R. Phoenix, Arizona.
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in Big Bend National Park, Texas.
Ph.D.
Dissertation, Univ. Arizona, Tucson. 172 p.
_________ , and P. R. Krausman.
1983.
Plant
communities of the lower desert shrubland in
Big Bend National Park,
Texas.
Second
Chihuahuan Desert Symposium, Alpine, Texas.
In Press.
Morista, M.
1959.
Measuring of interspecific
association
and
similarity
between
communities.
Mem. Fac. Sci. Kyrishu Univ.,
Sere E. BioI. 3:65-80.
The annual behavioral
Phillips, J. L.
1974.
cycle
of
desert mule deer
(Odocoileus
hemionus crooki) in relation to vegetative
use.
M.S. Thesis, SuI Ross State Univ.,
Alpine, Texas. 60 p.
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deer, Odocoileus hemionus crooki Mearns in
southeastern Arizona.
Ph.D. Dissertation,
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North
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Univ.
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H.
1974.
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Prentice-Hall Inc., Englewood Cliffs, N.J.
620 p.
Dix, R. L.
1969.
An application of the pointcentered quarter method of the sampling of
grassland vegetation.
J.
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14:63-69.
Johnson, R. R., S. W. Carothers, and J. M.
Simpson.
1981.
A riparian classification
system. p 375-382 in R. E. Warner, and K. M.
Hendrix, eds.
California riparian systems:
ecology,
conservation,
and
productive
management. Univ.
of
California
Press,
Berkeley. 1035 p.
Koerth, B. H., Jr.
1981.
Habitat use, herd
ecology, and seasonal movements of mule deer
in the Texas Panhandle.
M.S. Thesis, Texas
Tech Univ., Lubbock. 103 p.
Krausman, P. R., and E. D. Ables. 1981. Ecology
of the Carmen Mountains white-tailed deer.
National Park Service Sci. Monogr. 15:1-114.
__________ , J. J. Hervert, and L. L. Ordway.
1984.
Radio tracking desert mule deer and
bighorn sheep with light aircraft.
pp. 115118 in P. R. Krausman and N. S. Smith, eds.
Deer in the southwest: a workshop. School of
Renewable
Natural
Resources,
Univ.
of
Arizona, Tucson, Arizona.
1978.
Forage
relationships
between two deer species in Big Bend National
Park, Texas. J. Wildl. Manage. 42:101-107.
__________ ,
1984. Impacts of the Central Arizona
Project on desert mule deer and
desert
149