Biodiversity and management of the madrean archipelago

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One Hundred Years of Vicissitude:
Terrestrial Bird and Mammal Distribution
Changes in the American Southwest,
1890 -1990
1 2
David E. Brown and Russell Davis
Abstract.-Distribution changes over the past 100 years are
summarized for a number of species of terrestrial mammals and birds in
Arizona and New Mexico. At least 39 species appear to have been
extirpated or suffered range restrictions, while 55 others have
experienced range expansions. Even when exotics are excluded, the
biodiversity of endotherms is now greater than in 1890. As expected, the
ranks of the "losers" contain a disproportionate number of large
predators and grassland-associated animals. By way of contrast, the
majority of the "winners" were forest and/or scrubland-adapted species.
More germane to this study was the fact that more than 70 percent of
the "winners" were species which have their primary biotic affinity south
of the U.S.-Mexico border. Few Rocky Mountain, Great Basin, or Great
Plains species increased in distribution. Possible reasons for these
phenomena are discussed including the potential effect of increasing
winter temperature minima on plant distributions and the availability of
invertebrates.
..
.
Much well-deserved concern has recently been THE STUDY AREA
expressed regarding an increase in the number of
endangered species and a loss in biodiversity. The states of New Mexico and Arizona in the
Each year sees additional animals proposed for American Southwest provide an ideal study area
state or federal listing as "threatened" or "endan for a long term analysis of the effects of climatic
gered" species. But, although much effort is change on endothermic animals. The region's
expended to determine the actual status of these southern position astride the Continental Divide,
animals, the stated reasons for their decline are and its great range in elevations from less than 30
often vague and subjective. Rarely, and only re m where the Colorado River enters Mexico to
cently, have overall changes in the status of an more than 4,000 m atop Wheeler Peak in the San
animal's habitat and its associated biota been gre de Cristo Mountains, provide climates
properly evaluated and appreciated. To achieve a ranging, from subtropical to arctic-boreal. Also,
better perspective of the possible influence of an depending on location, mean annual precipitation
overall environmental change, we examined the can be as low as 75 mm to greater than 780 mm.
distributional status of terrestrial mammals and The result is that almost every biome found in
birds in the states of Arizona and New Mexico North America - alpine tundra, boreal conifer
during the past 100 years. forest, temperate deciduous forest, evergreen
woodland, chaparral, grassland and desert - can
be found somewhere in these two states. More
over, the Southwest's othography provides both
mountain and valley corridors extending north
1 Department of Zoology, Arizona State University. ward into the Rocky Mountains and Great Basin
as well as southward into Mexico. Thus, birds,
2Department of Ecology and Evolutionary Biology, University of Ari
zona. and many mammals, can? in a single day, travel
231
from an environment in which freezing tempera Another factor contributing to our under
tures may occur at any time of the year to those standing of the Southwest's biota was its
which rarely experiences a 24-hour freeze. relatively late settlement by Western Man. Ac
The American Southwest's biotic diversity is quired by the United Stated from Mexico in the
also due to its special evolutionary history. Most mid-19th Century, New Mexico Territory had re
of the region was not covered by ice sheets during mained virtually uninhabited by either Spaniards
the Pleistocene Epoch. Although the alternating or Mexicans. This circumstance enabled American
drying and cooling that accompanied glacial ad scientists to report on the region's landscapes and
vances and retreats had an immense effect on biota prim to large scale ranching and agricultural
Southwest environments, the region's great topo development (see e.g., Davis 1982). Moreover, this
graphic relief, manifested in such features as the late settlement, combined with the region's rug
Grand Canyon and Mogollon Rim, coupled with ged topography and general aridity, prolonged
it southern latitudes, served to provide warm the settlement process. Even as late as 1880, few
temperate refugia for a number of endemic settlements existed outside of the river valleys
Pleistocene plants and animals. These features, and the environs of mining camps. The late entry
coupled with the erratic but overall drying and of Arizona and New Mexico into the Union in
warming trend that accompanied the onset of 1912 also allowed for the inclusion of sizable
Holocene times about 11,500 years ago, have re tracts of land within National forests and parks
sulted in the Southwest being the location of before they were irreversibly impacted by a long
isolated Pleistocene forests on mountain-tops, rel history of unregulated grazing, land clearing and
ict mild-winter conifers and water-dependent other ecologically disruptive influences.
decid uous forests beneath the Mogollon Rim, and Last, but not least, the American Southwest
a tension zone of diminishing areas of grasslands was visited by a number of competent naturalists
and evergreen woodlands between 1000 and 2000 around the tum of the Century. People like Elliot
m elevation. The most recent phenomenon has Coues, E.A. Mearns, Herbert Brown, H.S. Swarth,
been the development and advance of the four Vernon Bailey and Florence Merriam Bailey made
North American deserts, parts of all of which oc systematic attempts to inventory the birds and
cur in the Southwest (see e.g., Van Devender et al. mammals of Arizona and New Mexico and re
1987). The result today is a great mixing of old ported on their habitat affinities and status (e.g.,
and new plant and animal species that have segre Swarth 1914). Other investigators conducted stud
gated out into a mosaic of Tertiary, Nearctic and ies on the distributions of particular species of
Neotropical habitats (Lowe and Brown~982). birds and mammals as well as the natural history
This biodiversity, coupled with the region's of specific locales (e.g., Merriam 1890). These
general aridity, has made the Southwest a focal studies have continued to the present day, allow
point of study for a variety of environmental dis ing for a progression of comprehensive treatments
ciplines. During the last 100 years the Southwest on the region's mammals and birds (e.g., Coues
has provi<;ied a unique field laboratory for zoolo 1867, Mearns 1907, Ligon 1927, Bailey 1928, Bailey
gists, botanists, ecologists, archeologists and 1931, Hall 1946, Ligon 1961, Phillips et al. 1964,
paleontologists. Not only has the region's abun Findley et al. 1975, Hubbard 1978, Monson and
dant fossil record been much investigated (see Phillips 1981, Hoffmeister 1986). Also relevant to
e.g., Harris 1985), much of the pioneering and on this study is the fact that 100 years of climatologi
going work on fossil pollen analysis, tree-ring cal data are now available for a number of New
chronology and other techniques to measure cli Mexico and Arizona stations (see e.g., Karl et al.
matic change have been focused in or near the 1989, Sellers et al. 1985).
Southwest. Especially germane to this effort is the
work of Hastings and Turner (1965) and their suc
cessors who have demonstrated important STUDY METHODS AND LIMITATIONS
landscape changes over the past 100 years
through the use of repeat-photography. The result Using as base references the basic works on
is that few if any regions of North America have a avian and mammalian distribution noted above,
more complete fossil and recent history of faunal state endangered animal summaries (Arizona
and vegetative changes. That these changes have Game and Fish Department 1988, New Mexico
been dramatic and have been shown to have con Department of Game and Fish 1992), specific jour
tinued into modern times, both facilitates and nal articles (e.g. Hock 1952, Hubbard 1977,
lends impetus to the work at hand. Lomolino et al. 1989) and our own work (e.g.
232
Brown 1973 and 1989, Davis and Dunford 1987, Rail (Rallus longirostris), Mexican Duck (Anas
Davis and Brown 1989, Davis and Callahan 1992), diazl), Black-bellied Whistling Duck (Dendrocy
we listed those species of terrestrial mammals and gna autumnaHs) and Cattle Egret (Bubulcus ibis)
birds which appear to have increased or de now enjoy a greater distribution in these two
creased their Southwest distributions during states than they did in 1900.
historic times. For comparative purposes we have There are, of course, a number of problems
also listed terrestrial species of exotics which are inherent in any comparative analysis of patterns
deemed to have established self-sustaining popu of past and present animal distributions. Perhaps
lations. To determine if the distribution of animals the foremost of these is that any knowl edge of the
in certain habitats had changed more than in other actual previous (a nd present) distribution of a
habitats, each species was assigned the biotic species is imperfect and open to interpretaiton.
community or communities with which it is pri The age-old question of whether a new locale for
marily associated. These lists were then a species represents a range extension or merely a
commented on and added to by John Hubbard, lack or failure of previous sampling efforts will
Gale Monson and Barry Spicer. Dr. Hubbard and never be answered in every case to everyone's sat
Mr. Monson are recognized authorities on the isfaction. The distributions of some species,
status of birds in New Mexico and Arizona, and especially those at the edge of their range, are es
Dr. Hubbard and Mr. Spicer have been involved in pecially dynamic and the occurrence of these
the preparation of status reports on rare and en species in Arizona or New Mexico is inherently
dangered species for their respective state sporadic (e.g. the Ghost-faced Bat (Mormoops
agencies. Their enthusiastic assistance was a ma megc11ophylla) and Rose-throated Becard
jor contribution to this study. (Pacllyramphus aglaiae) in Arizona. These species
Bird names follow Hubbard (1978) and Mon are either omitted from our analysis or indicated
son and Phillips (1981). Mammal names follow with a question mark. Moreover, the status and
Jones, et. al. (1992). The biotic community affili distribution of some species such as the Long
ations are those discussed in Brown (1982, 1994) nosed Bat (Leptonycteris curasoae = sanbornl) is a
and mapped by Brown and Lowe (1980, 1990). matter of some debate (see e.g., Howell and Roth
Our lists only include those species of terres 1981, Cockrum 1991). For purposes of this analy
trial birds and mammals known to have bred in sis, we have excluded those species which are
the Southwest states of Arizona and New Mexico known to irregularly occur in the American
since 1890. Subspecies are not included,. This lim Southwest or whose past and present distribu
ited geographical perspective is due both to a lack tional status is considered questionable by
of comparable historical data for the Mexican biologists involved in their study. Since this is a
states of Baja California, Sonora and Chihuaha study of the dynamics of distribution, we have
and to the fact that we have less experience with considered only changes in the distribution of
conditions in these states as well as those in Cali species, not an increase or decrease in population
fornia, Nevada and Texas. Regrettably, sizes.
comparable historical data and experience are also
lacking for the Southwest's herptile and inverte
brate faunas. Such information could have
contributed greatly to the breadth of our analysis. RESULTS
The massive changes that have occurred in the
Southwest's riparian and other wetland habitats Nine species have been were extirpated in Ari
also precludes a comparable analysis of the re zona and New Mexico (Tab Ie 1). Not surprisingly,
gion's aquatic and wetland faunas. However, it is three of these are large carnivores which were sys
important to point out that the dire status of the tematically hunted as livestock predators. One
Southwest's native fishes and their extensive dis other, the black-footed ferret, may have been lost
placement and replacement by "exotics" has been as a result of prairie dog control efforts although
I
well documented (see e.g. Minckley and Deacon the animal's demise due to one or more intro
1991). It goes without saying that endothermic d uced pathogens is a distinct possibility. Two
animals such as the River Otter (Lutra canadensis) large game animals, the elk and bison, formerly
and Beaver (Castor canadensis) have experienced preSE!nt and subsequently extirpated, have since
similar displacements. Nonetheless, it should also been reintroduced, the former with marked suc
be noted that a number of wetland birds including cess. The remaining three extirpated species are
the Bald Eagle (Halieatus leucocephalus), Clapper grassland-associated birds.
233
Thirty-four species are still present but appear mammals and two birds can be considered as
to have experienced range reductions since 1890 Rocky Mountain species.
while 55 species now seem to enjoy an expanded The above distributional changes strongl)
distribution (Tables 2 and 3). Although human-as suggest recent changes in both landscape charac
sisted introductions and the. occurrence of ter and climate. Indeed, a change in vegetatior
artificial feeding and watering sites can explain from open grasslands and woodlands to morf
some of the increases in distribution, the reasons closed communities of forest and scrubland has
for most are unclear and cannot be explained by been well documented for Arizona by Hastings
direct human intervention. Conversely, intro and Turner (1965) and in New Mexico by Dick
duced diseases, competition with exotic animals Peddie (1993). While climatic change has beer.
and grazing by livestock have been implicated as going on in the Southwest since the advent of the
reasons for many of the species now having de Holocene (see e.g., Houghton et al. 1990, Davis
creased distributions. and Shafer 1992), the amount of this occurring
Only 19 exotics (13 mammals and six birds) within a time frame of only 100 years is difficult to
appear to have become established in Arizona and assess. Climatic data since 1890 show no apprecia- .
New Mexico by 1890 (Table 4). The two camels ble trends in either summer or winter
have since been eliminated and the status of sev precipitation amounts in Arizona (Sellers et al.
eral other species (e.g., Swine, Chukar Partridge 1987) and changes in minimum temperatures are
and Black Rat) may be tenuous. A surprising re often masked by the effects of urban warming
sult of the analysis was that even when exotic (Kirby and Sellers 1987, Sellers 1990). Nonethe
animals are excluded and extirpated species in less, regional climatic data which has been
cluded, more animals appear to have experienced corrected for the effects of urban warming shows
range expansions (55) than declines (43). As far as a significant increase in winter, summer and an
terrestrial birds and mammals are concerned, bio nual temperature minima since ca. 1960 for the
diversity is now greater in the American "Southern Deserts" which encompasses southern
Southwest than it was in 1890. New Mexico and Arizona (Karl et al. 1989). No
Tables 5 and 6 summarize the formation-types, such trend in apparent for the "Great Basin" and
biotic community affinities and centers of bio "Southern Rockies" regions. Clearly, further in
geographic distributions of 39 mammals and birds vestigation into the possible effects of short-term
which appear to now have reduced distributions climatic change are warranted. It may be instruc
in the American Southwest. The majority (59%) of tive to moniter and evaluate short-term (50 to 100
these are grassland-affiliated species. "Nearly all of year) changes in the distributions of select species
the biogeographic provinces are represented, with of terrestrial ectotherms (invertebrates and herp
nearly as many species having their centers of dis tiles) to determine what role, if any, climate
tribution to the north or northeast (49% as to the change might also be playing on their distribu
)
south or southeast (51 %). tions in the American Southwest.
Formation-class affinities and biogeographic
centers for the 55 species of mammals and birds
breeding in the American Southwest and which LITERATURE CITED
have experienced range expansions are presented
in Tables 7 and 8. Twenty-nine, or more than half Arizona Game and Fish Department. 1988. Threatened
(53%), of these "winners" are forest and woodland native wildlife in Arizona. Arizona Game and Fish
species. Another seven (13%) are associated with Department, Phoenix.
chaparral or other scrub lands, only seven are Bailey, F.M. 1928. Birds of New Mexico. New Mexico
Department ofG ame and Fish. Santa Fe.
grassland-associated species and, oddly enough,
Bailey, V.1931.MammalsofNewMexico. USDAN.Amer.
only three (50/0) are desert species. The most sur
Fauna53:1-412.
prising statistic, however, is the number of species
Brown, D.E. 1973. Western range extensions of scaled
having their biogeographic center of distribution
quail, Montezuma quail and coppery-tailed trogon in
to the south of the American Southwest (Table 8).
Arizona. West. Birds 4:59-60.
More than half of the "winners" have their nearest Brown, D.E.1989. Arizona game birds. University of Ari
ecological center in either Mexico's Sierra Madre zona Press, Tucson.
(27%) or the forests and scrublands of Sonora and Brown, D.E. ed. 1982, 1994. Biotic communities -South
Sinaloa (29%). None of the "winners" are western United States and Northwestern Mexico.
headquarted in the Great Basin and only two University of Utah Press, Salt Lake City.
234
Brown,D.E.andC.H.Lowe,Jr.1980, 1990. Biotic communi American mammals north of Mexico. Occasional pa
ties of the Southwest (map). USDA Forest Service and pers. The Museum, Texas Tech University. No. 146, 23
U niversi ty of Utah Press, Sal t Lake City. pp.
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populations of the long-nosed bats, Leptonycteris san of regional season averages of maximum, minimum
borni Family Phyllostomidae. Anales Inst. BioI. Univ. and average temperature and diurnal temperature
Nac.Auton.Mexico, Ser. Zoo1.62:181-202. range across the United States: 1901-1987. Nation. Oce
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1:281-292,351-363,393-400,531-541. Nation. Climat. Data Cen., Ashevill, NC
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235
Table 1.-Nine species of terrestrial mammals and birds breeding in the American Southwest in 1890 and which were extirpated prior
to 1990.
Species Primary Biotic Affinities In the Southwest
MAMMALS
1
Ursus arctos, Grizzly Bear Rocky Mountain Subalpine Forests and Grasslands downward to Madrean
Evergreen Woodland and Interior Chaparral
Canis lUpus, Gray Wolf1 Rocky Mountain Montane Forests downward through Plains Grassland and
Madrean Evergreen Woodland to Chihuahuan Semidesert Grassland
Mustela nigripes, Black-footed Ferret2, 3 Great Basin Grassland
Panthera onca, Jaguar1 Madrean Evergreen Woodland and Riparian communities upward to Rocky
Mountain Conifer Forest
Cervus elaphus, Elk4 Rocky Mountain Conifer Forest and Meadow Grassland
4
Bos bison, American Bison Plains Grasslands in West Texas and New Mexico
IUB.Im
Falco femoralis, Aplomado Falcon Chihuahuan Semidesert Grassland
Centrocercus urophasianus, Sage Grouses Great Basin Desertscrub with Wet Meadows
pedjocetes phssjsnellus Sharp-tailed Grouses Intermountain Grassland interspersed with Scrub
1 Purposefully extirpated through direct human intervention,
2 Inadvertently extirpated through direct human intervention,
3 Shown to be susceptible to diseases transmitted by introduced animals.
4 Successfully reintroduced after 1912.
S Populations shown to be susceptible to livestock grazing and changes in land use.
236
Table 2.-Thirty-four species of terrestrial mammals and birds breeding In the American Southwest and experiencing range
reductions since 1890.
Species Primary Biotic Affinities in the Southwest
MAMMALS
Lepus alieni, Antelope Jackrabbit Sonoran Savanna Grassland, Sonoran Desertscrub, Chihuahuan Semidesert
Grassland
?Spermophi/us tridecemliniatus,
Thirteen-lined Ground Squirrel Plains and Great Basin Grasslands
Cynomys ludovicianus, Black-tailed Prairie Dog 1 Plains and Chihuahuan Semidesert Grasslands
Cynomys gunnisoni, Gunnison's Prairie Dog1 Great Basin Grassland
?Tamius minimus, Least Chipmunk Rocky Mountain Subalpine and Montane Conifer Forests and Montane Meadows
?Crategeomys castanops, Yellow-faced Pocket Gopher Plains Grassland
Microtus pennsylvanicus, Meadow Vole2 Rocky Mountain and Great Basin Meadows and Marshlands
?Sigmodon arizonae, Arizona Cotton Rat Grassy areas within Sonoran Desertscrub and various Riparian Biomes
Zapus hudsonius, Meadow Jumping Mouse2 Rocky Mountain Subalpine and Montane Meadows; Marshlands
1
VUlpes ve/ox, Kit Fox, Swift Fox ,3 Sonoran, Mohave and Chihuahuan Desertscrubs;Plains Grassland
1
Ursus americanus, Black Bear Rocky Mountain Subalpine and Montane Conifer Forests
Odocoi/eus virginianus, White-tailed Deer Madrean Evergreen Woodland, Chihuahuan Semidesert Grassland and
Riparian Forest within Interior Chaparral and other Biomes
Anti/ocapra americana, Pronghorn Plains and Chihuahuan Semidesert Grasslands: Sonoran Desertscrub
(Lower Colorado River Valley subdivision)
Ovis canadensis, Bighorn Sheep3 Sonoran and Mohave Desertscrub; Chihuahuan Semidesert Grassland;
various Great Basin and Rocky Mountain Biomes
8lB.tm
Buteo regalis. Ferruginous Hawk Great Basin Grassland
Buteo nitidus, Gray Hawk Sonoran and adjacent warm-temperate Riparian Woodlands of Mesquite and
Cottonwood trees
Caracara cheriway. Caracara Sonoran Desertscrub
Me/eagris gallopavo, Wild Turkey3,4 Madrean and Rocky Mountain Conifer Forests, Riparian Deciduous Forest
Tympanuchus pallidicinctus, Lesser Prairie-chickens Plains Grassland
Lagopus leucurus, White-tailed PtarmiganS Rocky Mountain Alpine Tundra and Shrubland
Cal/ipepla squamata, Scaled Quail Chihuahuan Semidesert Grassland
Colinus virginianus, BobwhiteS Plains Grassland; Sonora Savanna Grassland
Cyrtonyx montezumae, Montezuma QuailS Madrean Evergreen Woodland Savanna
Columbine passerina. Ground Dove Weedy fields within Chihuahuan Semidesert Grassland and Sonoran Desertscrub
Melanerpes erythrocephalus, Red-headed WoodpeckerS Riparian Deciduous Forest and telephone poles and trees within Plains Grassland
Glaucidlum brasilianum, Ferruginous Pygmy Owl Sonoran Desertscrub ~Arizona upland subdivision)
Tyrannus tyrannus, Eastern Kingbird Riparian Deciduous Forest and other Wooded Habitats within Plains Grassland
Empidonax fulvifrons, Buff-breasted Flycatcher Madrean Evergreen Woodland and Chaparral
?Vireo vicinior, Gray Vireo Great Basin Conifer Woodland, Interior Chaparral
?Passerina ciris, Painted Bunting Riparian Deciduous Woodlands
Ammodramus savannarum, Grasshopper Sparrow Plains Grassland, Chihuahuan Semidesert Grassland
Aimophila carpalis, Rufous-winged Sparrow Sonoran Savanna Grassland
Aimophila botterii, Botteri's Sparrow Chihuahuan Semidesert Grassland
Coccothraustes vespertinus EveningGrosbeak Rocky Mountain Subalpine Conifer Forest
I
1 Some isolated populations extirpated through direct human intervention.
2 Largely a wetland species in the Southwest.
3 Shown to be susceptible to diseases transmitted by introduced animals.
4 Also successfully introduced to non-native habitats.
S Populations shown to be susceptible to livestock grazing and/or browsing.
237
Table 3.-Fifty-five species of terrestrial mammals and birds that appear to have increased their breeding distribution In the
American Southwest since 1890.
Species Primary Biotic Affinities in the Southwest
MAMMALS
Didelphis virginian a , Virginia Opossum Various warm temperate Biomes in Southwestern Arizona
?Mormoops megalophylla, Ghost-faced Bat Riparian communities within Madrean Evergreen in Southwest Texas
Idionycteris phyllotis, Allen's Big-eared Bat Various warm and cold temperate Biomes
Nyctinomops femorosaccus, Pocketed Free-tailed Bat Sonoran Desertscrub and adjacent warm temperate Biomes
Dasypus novemcinctus, Nine-banded Armadillo Warm temperate Riparian communities along the Pecos River in West Texas
Sciurus aberti, Albert's Squirrel2 Madrean and Rocky Mountain Montane Conifer Forests
Baiomys taylori, Northern Pygmy Mouse Plains and Chihuahuan Semidesert Grasslands
?Sigmodon hispidus. Hispid Cotton Rat Chihuahuan Semidesert Grassland and various Riparian communities
?Sigmodon fulviventer, Tawny-bellied Cotton Rat3 Madrean Evergreen Woodland and Chihuahuan Semidesert Grassland
Sigmodon ochrognathus, Yellow-nosed Cotton Rat Madrean Evergreen Woodland Savanna and Montane Meadow Grassland
Microtus mexicanus, Mexican Vole Meadow and Grassland areas within Madrean, Rocky Mountain and
Great Basin Montane Biomes
Canis latrans, Coyote All, or nearly all, Biomes
?Vulpes vulpes, Red Fox Rocky Mountain Alpine Tundra and Subalpine Conifer Forest; may now be
extending its range into Plains and Great Basin Grasslands and other Biomes
Nasua narica, White-nosed Coati Madrean Evergreen Woodland
Gonepatus meso/eucus, Common Hog-nosed Skunk Various warm temperate Biomes
Tayassu tajacu. Collared Peccary Sonoran Desertscrub and adjacent warm temperate Biomes
Gervus elaphus, Elk4 Rocky Mountain Montane Forest and adjacent Meadows
4
Bos bison, Bison Plains and Great Basin Grasslands within and outside of historical range
Odocoileus hemionus, Mule Deer Nearly all Biomes having sufficient cover and not occupied by O. virginianus
BIRDS
Coragyps atratus, Black Vulture Sonoran Desertscrub
/ctinia mississippiensis, Mississippi Kite Riparian Deciduous Forest within Sonoran Desertscrub
Elanus leucurus, White-tailed Kite Riparian Deciduous Forest within Chihuahuan Semidesert Grassland
Parabuteo unicinctus, Harris Hawk Sonoran Desertscrub (Arizona upland division); Mohave Desertscrub;
Mesquite and other dense vegetation within Chihuahuan Desertscrub
Scardafella inca, I nca Dove Urban areas within Sonoran Desertscrub
Zenaida asiatica, White-winged Dove Sonoran Desertscrub and adjacent warm-temperate Biomes including
residential areas
Gaprilmulgus ridgwayi. Buff-collared Nightj ar Riparian communities within Chihuahuan Semidesert Grassland and
Sonoran Desertscrub
?Ghaetura pelagica, Chimney Swift Urban and residential areas
?Calothorax lucifer, Lucifer Hummingbird1 Riparian communities within Madrean Evergreen Woodland Savanna
Galypte anna, Anna Hummingbird1 Urban areas within Sonoran Desertscrub
?Amazilia berylina, Berylline Hummingbird1 Riparian communities within Madrean Evergreen Woodland
Amazilia violiceps, Violet-crowned Hummingbird1 Riparian communities within Madrean Evergreen Woodland
Trogon elegans, Coppery-tailed Trogon Riparian communities within Madrean Evergreen Woodland
Euptilotus neoxenus, Eared Trogan Riparian communities within Madrean Evergreen Woodland
Dendrocopos pubescens, Downy Woodpecker Deciduous trees within Rocky Mountain Subalpine and
Montane Forests; Riparian Deciduous Forest
Tyrannus crassirostris, Thick-billed Kingbird Riparian Deciduous Forests within Chihuahuan Semidesert Grassland
and Sonoran Desertscrub
Tyrannus meiancholicus, Tropical Kingbird Riparian deciduous w()odlands within Chihuahuan Semidesert Grassland
and Sonoran Desertscrub
Gyanocitta cristata. Blue Jay Riparian deciduous trees and residential areas
Parus atricapillus. Black-capped Chickadee Rocky Mountain Subalpine and Montane Conifer Forests: cold temperate
Riparian Deciduous Forest
Toxostoma bendirei, Bendire Thrasher Sonoran Desertscrub, Chihuahuan Desertscrub, Chihuahuan
Semidesert Grassland
Sialia sia/is, Eastern Bluebird Riparian Deciduous Woodland, Madrean Evergreen Woodland
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Heliminthophila luciae, Lucy Warbler Mesquite communities within Sonoran Desertscrub and various warm
temperate Riparian Biomes
Cardellina rubrifrons, Red-faced Warbler Madrean Evergreen Forest and Woodland
Cardinalis cardinalis, Cardinal Riparian communities within Sonoran Desertscrub (Arizona upland subdivision)
Cardinalis sinuatus, Pyrrhuloxia Denser communities within Sonoran and Chihuahuan Desertscrub
Piranga bidentata, Flame-colored Tanager Riparian communities within Madrean Evergreen Woodland
Icterus spurius, Orchard Oriole Riparian Deciduous Woodlands and residential areas in Southeast New Mexico
Icterus cucullatus, Hooded Oriole Tropical and warm temperate Riparian Deciduous Forests within Sonoran,
Chihuahuan and Mohave Biomes
Quiscaius mexicana, Great-tailed Grackle Urban and cultivated lands within Sonoran Desertscrub and various warm
temperate and cold temperate Biomes
Quisca/us quiscula, Common Grackle Urban and cultivated lands within various cold temperate Biomes
Mo/othrus ater, Brown-headed Cowbird Ranch yards, irrigated areas and Riparian areas from tropical to cold
temperate Biomes
Mo/othrus aeneus, Bronzed Cowbird Ranch yards, irrigated areas and Riparian communities within Chihuahuan
semidesert Grassland and Sonoran Desertscrub
Passerina cyanea, Indigo Bunting Riparian Woodlands within Sonoran Desertscrub as well as various warm and
cold temperate Biomes
Passerina versicolor, Varied Bunting Woody Riparian thickets within Sonoran Desertscrub and various
warm-temperate Biomes
Aimophila cassinii, Cassin Sparrow Chihuahuan Semidesert Grassland
Aimophila quinquestriata, Five-striped Sparrow Chihuahuan Semidesert Grassland
1 Range expansion possibly influenced by the provision of feeding stations.
2 Some, but not all, range expansions are due to introductions.
3 Despite what appears to be a general increase in distribution, at least one race (S. f. goldmam) of this species is believed to have been extir
fated in the Southwest.
Populations of this species have continued to increase in distribution since being reintroduced (and in some cases introduced) after 1912.
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Table 4.-Exotic species of terrestrial mammals and birds which became established In the American Southwest between 1890 and
1990.
Species Primary Biotic Affinities in the Southwest
MAMMALS
Scirus niger, Eastern Fox Squirrel Riparian Deciduous Forests, Orchards and residential Parks within
Chihuahuan and Sonoran Biomes
Rattus rattus, Black Rat1 In and around human habitations within Chihuahuan Semidesert Grassland,
Chihuahuan Desertscrub and Sonoran Desertscrub
Rattus norvegicus. Norway Rat In and around human habitations within a variety of Biomes
Mus musculus, House Mouse In and around human habitations; Chihuahuan Semidesert Grassland, Plains
Grassland and Riparian communities within several biomes
Came/ops spp., Dromedary and Bactrian Camels Mohave and Sonoran desertscrubs
Equus asinus, Feral Ass Sonoran and Mohave desertscrubs; Chihuahuan SemidesertGrassland,
Great Basin Desertscrub and Great Basin Conifer Woodland
Equus caballus, Feral Horse Chihuahuan Semidesert Grassland, Sonoran Desertscrub, Interior Chaparral,
Great Basin Conifer Woodland and possible other Biomes
Ammotragus lervia, Barbary Sheep1 Rugged areas within Plains, Chihuahuan and Great Basin
Oryx gazella, Gemsbok Chihuahuan Semidesert Grassland and Chihuahuan Desertscrub
Capra hircus, Goat1 Sonoran Desertscrub, Chihuahuan Semidesert Grassland, Interior Chaparral
Capra ibex, Ibex Interior Chaparral, Chihuahuan Semidesert Grassland, Great Basin Scrubland
and possibly other Biomes
Sus scrofa, Feral Pig Madrean Evergreen Woodland
BIRDS
A/ectorfs chukar, Chukar Great Basin Desertscrub and Grassland
Phasianus colchicus, Pheasant Agricultural areas within various warm temperate and cold temperate Biomes
Lophortyx californicus, California Quail Riparian Scrubland within Plains Grassland
Columba !ivia, Rock Dove In and around human habitations within various Biomes
Sturn us vulgaris, Starling In and around human habitations within various Biomes
Passer domesticus, House Sparrow In and around human habitations within various Biomes
1 Present status unknown; may now be absent from the Southwest.
2 Extirpated from the Southwest through direct human intervention by 1900.
3 Extirpated from native Plains Grassland in the South~est prior to 1890.
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