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Flora of the Huachuca Mountains,
Cochise County, Arizona
Janice E. Bowers and Steven P. Mclaughlin 1
Abstract.-The Huachuca Mountains, Cochise County, Arizona, are one
of about two dozen "sky islands" in southeastern Arizona. A herbarium
search revealed that, prior to 1990, 849 species had been documented
from the Huachuca Mountains. Field work conducted between 1990 and
1994 added another 144 species to the flora. Altogether, 993 species in
467 genera and 101 families are now known from the range. Of these,
65 are introduced. Madrean floristic elements dominate the flora,
accounting for 69.9% of all native species. Sonoran elements (5.0% of
all native species) are relatively poorly represented in the Huachuca
Mountains compared to more arid mountain ranges in southeastern
Arizona. The flora of the Huachuca Mountains is comparatively rich for
an Arizona local flora, with 29-39% more species than expected based
on its elevational range and collecting history. Substrate complexity and
the presence of many well-watered canyon habitats and springs
contribute to the high species diversity.
INTRODUCTION
Our initial objective was, based on the work of
these many collectors, to assemble a plant checklist for the entire range so that we could detemine
if the flora was indeed, as Wallmo (1955) characterized it, "quite well known." Eventually, we also
becalne interested in how plant checklists grow
and shrink. In this paper, we compare the flora of
the Huachuca Mountains with floras of other sky
islands in southeastern Arizona, and demonstrate
that species composition of local floras is dynamic, subject to historical changes in climate,
land use, and other factors. The checklist will be
published at a later date.
The Huachuca Mountains (fig. 1), located in
southwestern Cochise County on the United
States-Mexico border, are one of two dozen mountain ranges in southeastern Arizona. Often
referred to as "sky islands" (Heald 1951), these
ranges form a floristically diverse archipelago that
has been of keen interest to botanists for more
than a century. The Huachuca Mountains in particular have a long and illustrious botanical
history. Plant collection dates back to the botanical
explorations of John Gill Lemmon and Sara Plummer Lemmon in 1882 (Crosswhite 1979) and has
continued until the present day (fig. 2). Floristic
work includes an enumeration of Timothy E. Wilcox and Marcus E. Jones collections (Britton and
Kearney 1894, Jones 1930) and checklists for Fort
Huachuca, Ramsey Canyon, Garden Canyon and
Coronado National Memorial (Goodding 1950a,
1950b; Pratt 1963; Toolin 1980; Yatskievych 198081; Ruffner and Johnson 1991; Parfitt and Christy
1992). Altogether, 84 collectors have taken more
than 4000 specimens from the range.
1 University
STUDY AREA
The north-south trending Huachuca Mountains belong to the Basin and Range Province
(Hunt 1967). Maximum elevations are 9,466 feet
(2885 m) on Miller Peak, 9220 feet (2810 m) on
Carr Peak, 8725 feet (2659 m) on Ramsey Peak,
and 8410 feet (2563 m) on Huachuca Peak. Several
major canyons with perennial reaches drain the
precipitous eastern slope and eventually flow into
the San Pedro River. The western slope, part of the
Santa Cruz River watershed, has only a few
streams with perennial reaches. Overall, the
of Arizona, Tucson, AZ.
135
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Figure 1.-Huachuca Mountains and vicinity. The ....vy blllell line shows the study area boundary. A, Location of the Huachuca
Mountains In Arizona; B, admlnlstrlltlw units In and near the Huachuca Mountains; C, major drainages and peaks of the
Huachuca Mountains.
Huachuca Mountains appear highly dissected,
with a large ratio of canyon to ridge habitat.
Our study area had an elevational range of
4466 feet (1361 m) and covered about 122 square
miles (31,600 ha). The northern and eastern
boundaries roughly followed the base of the
range, which varies from 5000-5200 feet (15241585 m) above sea level. The southern edge
coincided with the International Boundary. The
5500-foot (1676 m) contour approximated the
western boundary. We excluded most private
lands at the base of the range, with the exception
of the Ramsey Canyon Nature Preserve, Peterson
Ranch in Scotia Canyon, and aquatic habitats at
Beatty's Miller Canyon Orchard in Miller Canyon.
The lower elevations of Fort Huachuca Military
Reservation were also excluded from our study
area.
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Figure 2.-Plant collection by YMr, Huachuca Mountain.
(omitting all years In which fewer tMn 10 apeclmen.
we,. collected). Based on specimen. depo.... at the
University of Arizona herbarium
136
Topography and Geology
THE FLORA
The range is geologically diverse. Bolsa
Quartzite, the basal sedimentary unit, rests unconformably on Precambrian granite. On the eastern
slope, Paleozoic sedimentary rocks, mainly limestone but also some shales and siltstones, top the
Bolsa quartzite. On the western slope, sedimentary rocks of Cretaceous age, including
conglomerates and shales, interfinger with Triassic-Jurassic volcanic and sedimentary rocks (Keith
and Wilt 1978).
Plant Checklist
In winter 1990 and spring 1991, we searched
the University of Arizona herbarium (ARIZ) and
the herbarium at Fort Huachuca for specimens
from the Huachuca Mountains. We critically
evaluated all collections, and, if necessary, redetermined them. Starting in August 1990 and
continuing through June 1994, we made 41 trips
into the range, mostly during the April-October
growing season, and also in November and January. We attempted to sample every habitat
throughout the growing season with special emphasis on discontinuous habitats such as cattle
tanks, springs, peaks, and cliffs. Most of our effort
was concentrated along trails and roads.
Between 1882 and 1989, collectors documented
a total of 849 species in the Huachuca Mountains.
During the course of our project, we found 137
species that were new to the flora. Another 7 species were added by other collectors between 1990
and 1993. The total flora comprises 993 species
and infraspecific taxa in 467 genera and 101 families. Of these, 65 species are introduced. The
native flora comprises 906 species and 27 infraspecific taxa.
Climate
Weather stations are maintained at Fort
Huachuca at the northern end of the range and at
Coronado National Memorial at the southern end.
Annual precipitation at Fort Huachuca (4664 feet,
1422 m) is 14.6 inches (37.1 cm). About half falls in
July and August as high-intensity "monsoonal"
rains that originate as scattered convectional
thunderstorms triggered and enhanced by surface
heating and orographic effects. Winters at Fort
Huachuca are rather dry. December and January,
the wettest winter months, average 1.7~ inches
(4.5 cm) of precipitation. About 10% of winter precipitation falls as snow, which seldom stays on the
ground more than a day or two. At higher elevations, annual rainfall exceeds 25 inches (63.5 cm),
and snow can remain on the ground all winter.
Winter storms result from cyclonic storms and
frontal systems associated with large-scale low
pressure systems that typically originate off the
coast of California and Baja California. They are
less variable spatially and more variable temporally than summer storms (Sellers and Hill 1974).
Summers and winters at Fort Huachuca are
mild. The average January temperature is 46.3°F
(7.9°C), with average daily maximum and minimum temperatures of 58.4 and 34.2°F (14.7 and
1.2°C). Summer temperatures are moderated by
afternoon cloud cover. The average July temperature is 77.5°P (25.3°C), with daily maximum and
minimum temperatures of 88.6 and 66.4°P (31.4
and 19.1°C). At higher elevations, the average
January temperature is 400 P (4.4°C), and the average July temperature is 65°P (18.3°C) (Sellers and
Hill 1974)0
How Plant Checklists Grow
It is common for plant checklists to expand
over several decades of collecting. The Mount
Shasta, California, flora grew from 425 species
and infra specific taxa in 1940 to 525 in 1963
(Cooke 1940, 1941, 1949, 1963), an increase of
about 1 percent per year. The flora of Tumamoc
Hill, Tucson Mountains, Arizona, increased 0.6
percent per year between 1909 and 1985, from 238
to 346 species (Thornber 1909, Bowers and Turner
1985). The flora of Organ Pipe Cactus National
Monument grew from 522 species in 1980 (Bowers
1980) to 571 in 1992 (Pinkava et al. 1992), an increase of 0.8 percent per year. The flora of the
White Mountains, California, increased 3 percent
per year between 1973 and 1987, from 761 to 1078
species (Lloyd and Mitchell 1973, Morefield 1992).
The small yearly increment in each case suggests
that the initial floras were fairly complete. Linear
regression of percent increase against final size of
the flora suggests that, not surprisingly, the larger
the flora, the more difficult it is to collect completely (R2 = 0.95).
137
had originally been collected before 1909. Rondeau (1991) did not find 55 species that had been
collected in the Tucson Mountains between 1903
and 1988. In the Huachuca Mountains, 31 species
may no longer belong to the flora, despite concerted efforts to locate many of them. Inevitably,
collectors of local floras seldom if ever relocate all
the plant species documented from an area. Some
are simply overlooked. Mislabeled vouchers
might not have belonged to the flora in the first
place. This is a particular problem for collectors in
southeastern Arizona, where Lemmon's labels are
notoriously unreliable (Kearney and Peebles
1960). Matelea balbisii/ Spirodela polyrhiza/
Woodsia scopulina and several other species reported from the Huachuca Mountains might well
have been mislabeled as to location. Some species
are "lost" as a result of taxonomic recombination;
in our study area, Aquilegia longisissima/ Polygala piliophora and a few others might not prove
to be good species.
Of greater biological interest are species that
apparently no longer occur in an area. Some
plants lost from the Huachuca Mountains flora are
exotics that apparently failed to become established, including Coriandrum sativum/ Pastinaca
sativa/ Lonicera japonica, and several others. A
few native species may have been eliminated by
development of the lower reaches of Carr, Miller,
and Ramsey canyons. Odontrichum decompositum, for example, was last seen at "James's resort"
near the mouth of Ramsey Canyon (Jones 1930),
now an area of houses, gardens, and pastures.
Melica porteri "cannot exist in the presence of a
cow" (Goodding 1950a) and may have been eliminated by grazing. Eventually, proliferation of
exotic grasses such as Eragrostis lehmanniana and
E curvula might have deleterious effects on the
native flora. Both species dramatically decrease
the diversity and productivity of native grasses
(Cable 1971, Bock et al. 1986). Eragrostis lehmannianil was introduced into Cochise County in the
late 1940s; by 1951 it had spread onto Fort
Huachuca, apparently from nearby highways
(Goodding 1950a). Eragrostis curvula was introduced into the United States in 1928 (Crider 1945).
These two exotics are now among the most common plants at low to moderate elevations in the
Huachuca Mountains.
The fossil record demonstrates dramatic alteration of floras as a result of climatic change
(Betancourt et al. 1990). Historical climatic change
repeats this process on a briefer time-scale, especially among small, local populations. Extirpation
of 6 native perennials formerly found at high ele-
Most additions to local floras are probably
plants that have been previously overlooked. On
occasion, however, movement of species onto a
site increases the size of a local flora. New arrivals
may be natives or exotics. In either case, careful
observation is needed to distinguish newly arrived species from those that were simply
overlooked. Especially in recent years, introduction of exotics, either deliberately or accidentally,
has expanded the size of many local floras. At
Glacier National Park, Montana, exotics increased
at an accelerating rate between 1920 and 1993 (Lesica et al. 1993). Over 76 years, the number of
introduced species in the Tumamoc Hill flora increased by an order of magnitude, from 2 to 52
(Bowers and Turner 1985, Burgess et al. 1991).
Such examples could be multiplied many times.
In the Huachuca Mountains flora, the 65 exotics
include species seeded by the Forest Service to
prevent erosion after fire (Dactylis glomerata/
Sanguisorba mino!; Melilotus spp.); escapes from
cultivation (Pyracantha koidzumii Hedera helix/
Vinca majo!; Rubus procera); and naturalized exotics (Erodium cicutarium/ Bromus rubens/
Polypogon interruptus). About half of the exotic
flora was first documented after 1962. The rapid
vegetative growth and smothering habit of Rubus
procera and Vinca major represent serious threats
to the biodiversity of lower mesic canyons in the
Huachuca Mountains.
The concentrated effort of compiling a plant
checklist is another reason local floras increase in
size. After 98 years of casual and infrequent collecting in the Rincon Mountains, the documented
flora was 517 species (Bowers, unpublished data).
The final checklist of 986 species (Bowers and
McLaughlin 1987) represented an increase of 18
percent per year. McLaughlin (1993) doubled the
known size of the Pinalefto Mountains flora, from
406 in 1988 (Johnson 1988) to 824 in 1993. In the
Huachuca Mountains, collectors documented a total of 849 species between 1882 and 1989. Between
1990 and 1994, 144 species were added to the
flora, an increase of about 5 percent per year. This
modest increment suggests that the flora was indeed comparatively well known at the start of our
project.
How local Floras Shrink
Loss of species from local floras has been
noted infrequently in southeastern Arizona. Bowers and McLaughlin (1987) were unable to relocate
41 species in the Rincon Mountains; of these, 22
138
vations in the Rincon Mountains might have resulted from severe winter drought during
1920-1930 or 1942-1958 (Bowers and McLaughlin
1987). During the seasonally dry months of April,
May and June, such species depend on soil moisture left by the winter snowpack. Severe winter
drought could have eliminated their presumably
small populations. A similar set of circumstances
might have claimed some species in the Huachuca
Mountains. High-elevation mesophytes not collected there since 1913 include Adiantum
pedatum, Achillea mil1elolium, Dugaldia
hoopesii, Macromeria viridillora, Sidalcea
neomexicana, and Veronica serpyl1ilolia. Cur-.
rently, Vaccinium myrtillus, Pyrola chlorantha,
Mertensia Iranciscana, Pedicularis grayi, Actaea
rubra, Hypericum lormosllm, Senecio huachucanus and several others are known in the
Huachuca Mountains only from small populations at high elevations and may be similarly
vulnerable.
1
Natural disasters, particular 'fire and flood,
may also have eliminated species from the flora.
The Huachuca Mountains have experienced frequent severe fires in recent years (Taylor 1991,
Wohl and Pearthree 1991). The southern end of
the range has been particularly hard-hit, notably
in 1977, 1988, and 1991 (Taylor 1991). The disappearance or retreat of several species can perhaps
be ascribed to these or earlier fires. Rosa woodsii,
collected in "moist draws" on Carr Peak in 1909,
is now known only from upper Bear Canyon; Hypericum lormosum, collected on "moist slopes" of
Carr Peak in 1909, is now known only from Bond
and Sawmill springs. Valeriana edulis, which is no
longer known from the flora, may have been
eliminated by fire.
The most destructive fires may be followed by
floods and debris flows, especially in steep drainages (Wohl and Pearthree 1991), with dire
consequences for riparian herbs (Gori 1992). The
small, scattered populations of Lilium parryi have
experienced catastrophic declines in recent years
as a result of flooding (Warren and Reichenbacher
1991, Wood 1992). Riparian plants that might have
been eliminated from the Huachuca Mountains
flora by floods or debris flows are Dryopteris
filix-mas, Aster coerulescens, Monarda fistulosa,
Oenothera kunthiana, Rubus arizonensis and
Glyceria borealis. Small populations of riparian
plants might have been eliminated during
drought years, as well.
Clearly, a variety of natural and man-made
disasters can eliminate species from local floras,
particularly when populations are small and local.
139
Although our failure to relocate 31 species provides only negative evidence, we find it
suggestive that distinct patterns such as fire,
flood, drought and development can be identified. Some apparently extirpated species might
well still occur in less accessible parts of the
mountain range.
FLORISTIC ANALYSIS
All native species occurring in the Huachuca
Mountains flora were classified into floristic elements based on the system of floristic areas for the
western United States developed by McLaughlin
(1992). Methods for assigning species to floristic
elements are given in McLaughlin and Bowers
(1990) and McLaughlin (1994). For comparison,
the floristic analysis of the Huachuca Mountains
flora is presented along with those from the Rincon and Pinalefto mountains (Bowers and
McLaughlin 1987, McLaughlin 1993) (Table 1).
The system of floristic elements of McLaughlin (1992) is hierarchical. Five floristic provinces
are recognized for the western United States: Cordilleran, Intermountain, Sonoran, Californian and
Madrean. These provinces are subdivided into
subprovinces, which are in turn subdivided into
districts. Table 1 provides a breakdown of floristic
elements for the Madrean Floristic Province.
"Widespread" Madrean species are those that are
(1) found in 20 or more of the local floras used by
McLaughlin (1992) to develop the classification,
and (2) centered on the Madrean Floristic Province. "Regional" Madrean species are ·those with
more restricted distributions (found in 10 to 19 of
101 local floras from the western United States)
that are centered within the Madrean Floristic
Province. "Central Arizonan," "Chihuahuan," and
"Apachian" species are narrowly distributed in
Table 1.-Florlstlc elements In the Huachuca, Rincon and
Plnaleflo mountains. Values In the table are the
percentage of the total native flora assigned to each
floristic element.
Huachuca
EIQri§li~ Elemeo§
Madrean
Widespread
Regional
Central Arizonan
Chihuahuan
Apachian
Total Madrean
Sonoran
Cordilleran
Intermountain
~alifQ[[]ia[)
MQ!.mt~i[)§
Rincon
MQUOtAio§
Pinalerio
MQUOtAilJ§
5.6
17.8
1.3
6.4
5.1
19.0
1.5
4.3
6;6
20.5
2.3
2.4
~
at.6
2.M
69.9
61.5
52.7
5.0
18.0
3.5
19.4
10.6
2.9
10.7
27.2
4.6
36
56
~6
Huachuca Mountains, where winters are neither
as cold nor as wet, there is no spruce-fir forest,
and the mixed-conifer forest is of limited extent.
In the sky island region in general, and in the
Huachuca Mountains in particular, the Cordilleran elements include both many widespread taxa
and many narrowly dis~ributed species with Mogollon affinities (centered in the Mogollon Rim of
Arizona and the Mogollon highlands of New
Mexico). In the Huachuca Mountains, species
with Cordilleran and Mogollon affinities are
found mostly at high elevations or in moist,
shaded canyons.
the western United States (found in 9 or fewer of
the sample of 101 local floras) and are centered,
respectively, within the Central Arizona, Chihuahuan and Apachian floristic districts.
The Huachuca, Rincon and Pinaleno mountains all lie within the Madrean Floristic Province,
since the majority of their species belong to Madrean elements. All three floras are placed within
the Apachian district, since the Apachian element
is the largest narrow-species element in their respective floras. The Huachuca Mountains flora
has the highest percentage of Apachian species,
and all species with Madrean affinities constitute
nearly 70% of the flora. The Cordilleran element
accounts for 18% of the Huachuca Mountains
flora; species with Sonoran and Californian affinities are better represented in the flora of the
Rincon Mountains, and those with Cordilleran
and Intermountain affinities are better represented in the Pinaleno Mountains. The Chihuhuan
element is somewhat better represented in the
Huachuca Mountains than in the Rincon or Pinaleno mountains.
The importance of Madrean floristic elements
in the Huachuca Mountains is not unexpected.
The Apachian element is particularly large. Genera notably rich in Apachian species (5 or more) in
our study area include Asclepias/ Bidens/ Brickellia/ lpomoea/ Dalea/ Desmodium/ Cyperus, and
Muhlenbergia. The Apachian element is most
strongly associated with oak and pine-qak woodlands, plant communities that are particularly
well represented in the Huachuca Mountains.
The low percentage of Sonoran elements in the
Huachuca Mountains contrasts with that of the
Rincon Mountains. The base elevation of the Rincon Mountains is 2000 feet (610 m) lower than that
of the Huachuca Mountains, resulting in hotter
summers, milder winters and lower rainfall, all
conducive to a higher representation of species
with Sonoran affinities. Those Sonoran species
within the flora of the Huachuca Mountains are
mostly species with widespread and regional distributions. Some are spring-flowering annuals, a
group that is not well represented in the
Huachuca Mountains.
The Cordilleran elements are of much greater
importance in the Pinaleno Mountains, especially
above 9000 feet (2743 m) (McLaughlin 1993), than
in the Huachuca Mountains (Table 1). Species
with Cordilleran affinities are found mostly in the
mixed-conifer and spruce-fir forests in the Pinaleno Mountains, where moisture-loving,
cold-tolerant plants thrive under high winter precipitation and low winter temperatures. In the
SPECIES DIVERSITY
The plant species diversity of the Huachuca
Mountains was evaluated in two ways. First,
based on the elevational range of the study area
and the collecting effort invested in compiling its
flora, we compared the actual number of species
observed with the number expected to occur
(Bowers and McLaughlin 1982). The elevational
range of our study area is 4466 feet (1361 m). For
collecting time, we estimated low and high values
based on the number of years in which 50 or more
specimens were collected (22 years) and the
number of years in which 75 or more specimens
were collected (16 years). The results showed that,
compared to other local floras from throughout
Arizona, the Huachuca Mountains have 29-39%
more species than would be predicted based on
elevational range and collecting history.
We plotted number of species versus elevational range for the Huachuca Mountains and 23
other local floras from Arizona and New Mexico
(fig. 3). The regression line in figure 3 shows the
relationship between elevational range and richness in this sample of 24 floras [5 = 264 +
0.274(ilE), R2 = 0.502, P < .001)]. The flora for the
Huachuca Mountains is the farthest above the regression line, that is, it has the highest residual
value. Of the mountain ranges from the southwestern United States whose floras have been
investigated in detail, the Huachuca Mountains
appear to be exceptional in their high species diversity.
In local floras from the western United States,
elevational range is closely correlated with habitat
diversity, since both temperature and precipitation vary with elevation, often over short
distances. Habitat diversity in turn is a major determinant of species diversity. Thus local floras
from areas spanning a large elevational range
140
(such as the Huachuca Mountains) tend to have
higher species diversity (fig. 3). The Huachuca
Mountains also have many aquatic habitats and
much variation in gelogical substrates, and these
factors probably also contribute to high species
diversity in the range. The complex topography of
the range, with its numerous deep canyons cutting nearly to the ridge lines, makes a
topographically patchy landscape that may promote high species diversity. Bennett and
Kunzmann (1992) attempted to quantify topographic "roughness" and found that their index
was correlated with species diversity among a
small set of floras from the sky island region. In
some sky island floras, a pronounced biseasonal
rainfall regime allows both a spring and a summer flora to flourish (Bowers and McLaughlin
1987, McLaughlin and Bowers 1990). In the
Huachuca Mountains, where winters are rather
dry, spring-flowering species are not well represented.
Groombridge (1992) lists the" Apachian/ Madrean" region as one of 164 global centers of plant
diversity. Floras from the sky island region are
inherently richer than other floras from the western United States (McLaughlin, this symposium).
For example, the floras of the- Huachuca Mountains, Rincon Mountains, Sycamore Canyon,
Chiricahua National Monument and Buenos Aires
National Wildlife Refuge, all in the sky island region, are farthest above the regression line in
Figure 3. Much of the high species diversity of the
Huachuca Mountains is due to the presence of a
large Apachian component. Although it is clear
that certain floristic elements and floristic areas
are richer than others, the environmental, historical and ecological factors that determine these
inherent differences are not yet well understood.
1200
CONCLUSIONS
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HUACHUCAS
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After 112 years of plant collection, the flora of
the fIuachuca Mountains is well known. Local floras can never be completely collected. Additional
field work inevitably turns up species that had
been overlooked. Ornamental and crop plants invade from nearby settlements, sometimes
becoming naturalized. Native plants, too, may occupy new territory. Natural disasters such as fire,
flood, and drought may extirpate some species,
especially those with small populations in limited
habitats. Human-induced disasters such as grazing and plant introduction may also take a toll.
Inevitably, some species will be overlooked by
collectors and mistakenly assumed to be extirpated. Between 1882 and 1994, additions (144) to
the Huachuca Mountains greatly exceeded subtractions (31). Plant checklists serve as a baseline
for assessing floristic change in future decades.
In a region known for its biological diversity,
the I-Iuachuca Mountains are exceptionally rich in
plant species. Contributing factors include a large
Apachian floristic element, complex topography, a
wide elevational range, and a diversity of geological substrates and aquatic habitats.
800
W
~
z
RM
OA
" FB
400
"
• PF
200
SA;R
AM"·
TM
~M.
CC
.OP
" NHM
,,!M • AC
• WT
::l
z
•
• CW
• NM
0
0
500
1000
1500
2000
2500
ELEVATION RANGE (Meters)
figure 3.-Relationshlp between elevatlonal range (difference between highest and lowest elevations) and species diversity
among 24 local floras from Arizona and New Mexico. In addition to the Huachucas, the floras plotted are: AC, Aravalpa
Canyon, AZ (Warren and Anderson 1980); AM, Animas Mountains, NM (Wagner 1973); BA, Buenos Aires National Wildlife
Refuge, AZ (McLaughlin 1992b); CC, Canyon de Chelly National Monument, AZ (Halse 1973); CNM, Chirlcahua National
Monument, AZ (Reeves 1976); CR, Cooke's Range, NM (Columbus 1988); CW, Chiricahua Wilderness, AZ (Leith liter
1980); OM, Datil Mountains, NM (Fletcher 1972); FB, Fort
Bowie National Historic Site, AZ (Warren et al. 1992); MM,
Mule Mountains, AZ (Wentworth 1982); NHM, Northern Hualapai Mountains, AZ (Butterwick et al. 1991); NM, Navajo
National Monument, AZ (Brotherson et al. 1978)j NSR, Northern Santa Rita Mountains, AZ (McLaughlin and Bowers 1990);
Op, Organ Pipe Cactus National Monument, AZ (Bowers
1980); PF, Petrified Forest National Park, AZ (Petrified Forest
National Park 1976); PM, Pinaleno Mountains, AZ (McLaughlin
1993); RM, Rincon Mountains, AZ (Bowers and McLaughlin
1987); SC, Sycamore Canyon, AZ (Toolin et al. 1980); SA,
Sierra Ancha, AZ (Pase and Johnson 1968); TM, Tucson
Mountains, AZ (Rondeau 1991); WM, White Mountains, NM
(Hutchins 1974); WT, White Tank Mountain Regional Park, AZ
(Keil1973); WU, Wupatki National Monument, AZ (McDougall
1962).
LITERATURE CITED
Benn(~tt, P. S. and
M. R. Kunzmann.1992. Factors affecting
plant species richness in the Madrean Archipelago
north of Mexico. In A. M~ Barton and S. S. Sloane,
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141
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