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Cactus Ferruginous Pygmy-owl (Glaucidium brasilianum cactorum)
Background
The cactus ferruginous pygmy-owl (Glaucidium brasilianum cactorum)
(pygmy-owl) is in the order Strigiformes and the family Strigidae. It
is a small bird, approximately 17 centimeters (cm) (6.75 inches (in))
long. Males average 62 grams (g) (2.2 ounces (oz)), and females average
75 g (2.6 oz). The pygmy-owl is reddish brown overall, with a cream-
colored belly streaked with reddish brown. Color may vary, with some
individuals being more grayish brown. The crown is lightly streaked,
and a pair of black/dark brown spots outlined in white occur on the
nape suggesting ``eyes.'' This species lacks ear tufts, and the eyes
are yellow. The tail is relatively long for an owl and is colored
reddish brown with darker brown bars. The pygmy-owl is primarily
diurnal (active during daylight) with crepuscular (active at dawn and
dusk) tendencies. They can be heard making a long, monotonous series of
short, repetitive notes, mostly during the breeding season.
The pygmy-owl is one of four subspecies of the ferruginous pygmy-
owl. It occurs from lowland central Arizona south through western
Mexico to the States of Colima and Michoacan, and from southern Texas
south through the Mexican States of Tamaulipas and Nuevo Leon. Only the
Arizona population of the pygmy-owl is listed as an endangered species
(62 FR 10730; March 10, 1997).
The total number of pygmy-owls and their distribution in Arizona
are unknown. Survey and monitoring work in Arizona resulted in
documenting 41 adult pygmy-owls in 1999, 34 in 2000, 36 in 2001, and,
most recently, 18 in 2002. A cumulative total of 85 occupied sites
(includes both single or paired birds) were recorded during these 4
years (Abbate et al. 1999, 2000, AGFD unpubl. data). Most of these
pygmy-owls were distributed in four general areas: northwest Tucson,
southern Pinal County, Organ Pipe Cactus National Monument, and the
Altar Valley. We believe that more pygmy-owls exist in Arizona, but
systematic surveys have not been conducted in all areas of potential
habitat.
In addition, recent survey information has shown pygmy-owls to be
more numerous adjacent to and near the Arizona border in Mexico (Flesch
and Steidl 2000). There also exists considerable unsurveyed habitat on
the Tohono O'odham Nation, and, although we have no means of
quantifying this habitat, the distribution of recent sightings on non-
Tribal areas east, west, and south of the U.S. portion of the Tohono
O'odham Nation lead us to reasonably conclude that these Tribal lands
may support meaningful numbers of pygmy-owls. Consequently, we believe
that it is highly likely that the overall pygmy-owl population in
Arizona is maintained by the movement and dispersal of owls among
groups of pygmy-owls in southern Arizona and northern Mexico resulting
from the connectivity of suitable habitat. The extent to which pygmy-
owls disperse across the U.S./Mexico border is unknown. Therefore,
addressing habitat connectivity and the movements of pygmy-owls within
Arizona is the primary consideration of this proposal due to the
importance of maintaining dispersal and movement among pygmy-owl
groups.
Given recent data, it is probable that conservation of the pygmy-
owl in Arizona requires both sufficient numbers and productivity of
pygmy-owls north of the border and immigration of pygmy-owls from
Mexico into Arizona, although we do not know at this time to what
extent immigration does or needs to occur.
The patchy, dispersed nature of the pygmy-owl population in Arizona
suggests that the overall population may function as a metapopulation.
A metapopulation is a set of
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subpopulations within an area, where movement and exchange of
individuals among population segments is possible, but not routine. A
metapopulation's persistence depends on the combined dynamics of the
productivity of subpopulations, the maintenance of genetic diversity,
the availability of suitable habitat for maintenance and expansion of
subpopulations, and the ``rescue'' of subpopulations that have
experienced local extinctions by the subsequent recolonization of these
areas by dispersal from adjacent population segments (Hanski 1999,
Hanski and Gilpin 1991, 1997). The local groups of pygmy-owls within
Arizona may function as subpopulations within the context of
metapopulation theory. However, more information is needed regarding
the population dynamics of pygmy-owls in Arizona.
Historically, pygmy-owls were recorded in association with riparian
woodlands in central and southern Arizona (Bendire 1892, Gilman 1909,
Johnson et al. 1987). Plants present in these riparian communities
included cottonwood (Populus fremontii), willow (Salix spp.), ash
(Fraxinus velutina), and hackberry (Celtis spp.). However, recent
records have documented pygmy-owls in a variety of vegetation
communities such as riparian woodlands, mesquite (Prosopis velutina,
and P. glandulosa) bosques (Spanish for woodlands), Sonoran
desertscrub, semidesert grassland, and Sonoran savanna grassland
communities (see Brown 1994 for a description of these vegetation
communities). While native and nonnative plant species composition
differs among these communities, there are certain unifying
characteristics such as the presence of vegetation in fairly dense
thickets or woodlands, the presence of trees, saguaros (Carnegiea
giganteus), or organ pipe cactus (Stenocereus thurberi) large enough to
support cavities for nesting, and elevations below 1,200 meters (m)
(4,000 feet (ft)) (Swarth 1914, Karalus and Eckert 1974, Monson and
Phillips 1981, Johnsgard 1988, Enriquez-Rocha et al. 1993, Proudfoot
and Johnson 2000). Large trees provide canopy cover and cavities used
for nesting, while the density of mid- and lower-story vegetation
provides foraging habitat and protection from predators, and it
contributes to the occurrence of prey items (Wilcox et al. 2000).
The density of trees and the amount of canopy cover preferred by
pygmy-owls in Arizona has not been fully defined. However, preliminary
results from a habitat selection study indicate that nest sites tend to
have a higher degree of canopy cover and higher vegetation diversity
than random sites (Wilcox et al. 2000). Overall vegetation density may
not be as important as patches of dense vegetation with a developed
canopy layer interspersed with open areas. Vegetation structure may be
more important than species composition (Wilcox et al. 1999, Cartron et
al. 2000). This is related to the fact that canopy cover and layers of
vegetation provide hunting perches, thermal cover, and promote predator
avoidance regardless of species. Larger trees with greater canopy also
have a greater potential to support cavities needed for nesting. Flesch
(1999) indicated that areas with large trees and canopy coverage are
likely important areas for pygmy-owls in the Altar Valley. Riparian and
xeroriparian (dry washes) areas, which are often used by pygmy-owls,
are generally characterized by increased vegetation layers, higher
plant diversity and larger tree sizes because of increased moisture
availability.
Food
Pygmy-owls typically hunt from perches in trees with dense foliage
using a perch-and-wait strategy; therefore, sufficient cover must be
present within their home range for them to successfully hunt and
survive. Pygmy-owls also hunt by inspecting tree and saguaro cavities
for other nesting birds, and possibly bats. Their diverse diet includes
birds, lizards, insects, and small mammals (Bendire 1888, Sutton 1951,
Sprunt 1955, Earhart and Johnson 1970, Oberholser 1974, Proudfoot 1996,
Abbate et al. 1996,1999). Observations in Arizona from 1996 through
1998 indicate that reptiles, birds, mammals, and insects were 44, 23,
6, and 3 percent, respectively, of pygmy-owl prey deliveries recorded;
24 percent were unidentified (Abbate et al. 1999). It is likely that
use of insects was underestimated in these observations because of the
speed at which they are consumed and the difficulty in observing such
small prey items. The density of annual plants and grasses, as well as
shrubs, may be important to enhancing the pygmy-owl's prey base.
Vegetation communities which provide a diversity of structural
layers
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and plant species likely contribute to the availability of prey for
pygmy-owls (Wilcox et al. 2000). Pygmy-owls also utilize different
groups of prey species on a seasonal basis. For example, lizards, small
mammals, and insects are utilized as available during the spring and
summer during periods of warm temperatures (Abbate et al. 1999).
However, during winter months, when low temperatures reduce the
activity by these prey groups, pygmy-owls likely turn to birds as their
primary source of food and appear to expand their use area in response
to reduced prey availability (Proudfoot 1996). Therefore, conservation
of the pygmy-owl should include consideration of the habitat needs of
prey species, including structural and species diversity and seasonal
availability. Pygmy-owl habitat must provide sufficient prey base and
cover from which to hunt in an appropriate configuration and proximity
to nest and roost sites.
Water
Free-standing water does not appear to be necessary for the
survival of pygmy-owls. During many hours of research monitoring,
pygmy-owls have never been observed directly drinking water (Abbate et
al. 1999, AGFD unpubl. data). It is likely that pygmy-owls meet much of
their biological water requirements through the prey they consume.
However, the presence of water may provide related benefits to pygmy-
owls. The availability of water may contribute to improved vegetation
structure and diversity which improves cover availability. The presence
of water also likely attracts potential prey species improving prey
availability.
Reproduction and Rearing of Offspring
Male pygmy-owls establish territories using territorial-
advertisement calls to repel neighboring males and attract females.
Usually, pygmy-owls nest as yearlings (Abbate et al. 1999, Gryimek
1972), and both sexes breed annually thereafter. Territories normally
contain several potential nest-roost cavities from which responding
females select a nest. Hence, cavities/acre may be a fundamental
criteria for habitat selection. Historically, pygmy-owls in Arizona
used cavities in cottonwood, mesquite, ash trees, and saguaro cacti for
nest sites (Millsap and Johnson 1988). Recent information from Arizona
indicates nests were located in cavities in saguaro cacti for all but
two of the known nests documented from 1996 to 2002 (Abbate et al.
1996, 1999, 2000, AGFD unpubl. data). One nest in an ash tree and one
in a eucalyptus tree were the only non-saguaro nest sites (Abbate et
al. 2000).
Pygmy-owls exhibit a high degree of site fidelity once territories
(the area defended) and home ranges (the area used throughout the year)
have been established (AGFD unpubl. data). Therefore, it is important
that habitat characteristics within territories and home ranges be
maintained over time in order for them to remain suitable. This is
important for established owl sites, as well as new sites established
by dispersing pygmy-owls.
Shrubs and large trees also provide protection against predators
for juvenile and adult pygmy-owls and cover from which they may capture
prey (Wilcox et al. 2000). Little is known about the rate or causes of
mortality in pygmy-owls; however, they are susceptible to predation
from a wide variety of species. Documented and suspected pygmy-owl
predators include great horned owls (Bubo virginianus), Harris' hawks
(Parabuteo unicinctus), Cooper's hawks (Accipiter cooperii), screech-
owls (Otus kennicottii), and domestic cats (Felis catus) (Abbate et al.
2000, AGFD unpubl. data). Pygmy-owls may be particularly vulnerable to
predation and other threats during and shortly after fledging (Abbate
et al. 1999). Arizona Game and Fish Department (AGFD) telemetry
monitoring in 2002 indicated at least three of the nine young were
killed by predators prior to dispersal during a year when tree species
failed to leaf out due to drought conditions (AGFD unpubl. data).
Therefore, cover near nest sites may be important for young to fledge
successfully (Wilcox et al. 1999, Wilcox et al. 2000). A number of
fledgling pygmy-owls have perished after being impaled on cholla
cactus, probably due to undeveloped flight skills (Abbate et al. 1999).
Conditions which promote the proliferation of cholla (overgrazing,
vegetation disturbance, etc.) may contribute to this mortality factor.
Habitat that provides for successful reproduction and rearing of young
provides trees and cacti that are of adequate size to provide cavities
in proximity to foraging, roosting, sheltering and dispersal habitats,
in addition to adequate cover for protection from climatic elements and
predators in an appropriate configuration in relation to the nest site.
The primary constituent elements determined necessary for the
conservation of the pygmy-owl include: (1) Elevations below 1,200 m
(4,000 ft) within the biotic communities of Sonoran riparian deciduous
woodlands; Sonoran riparian scrubland; mesquite bosques; xeroriparian
communities; tree-lined drainages in semidesert, Sonoran savanna, and
mesquite grasslands; and the Arizona Upland and Lower Colorado River
subdivisions of Sonoran desertscrub (see Brown 1994 for a description
of vegetation communities); (2) nesting cavities located in trees
including, but not limited to cottonwood, willow, ash, mesquite, palo
verde, ironwood, and hackberry with a trunk diameter of 15 cm (6 in) or
greater measured 1.4 m (4.5 ft) from the ground, or large columnar
cactus such as saguaro or organ pipe greater than 2.4 m (8 ft); (3)
multilayered vegetation (presence of canopy, mid-story, and ground
cover) provided by trees and cacti in association with shrubs such as
acacia, prickly pear, desert hackberry, graythorn, etc., and ground
cover such as triangle-leaf bursage, burro weed, grasses, or annual
plants. By way of description, preliminary data gathered by AGFD
indicates 35 percent ground cover at perch sites and 48 percent ground
cover at nest sites; mid-story cover of 65 percent at perch sites and
65 percent at nest sites; and 73 percent canopy cover at perch sites
and 87 percent canopy cover at nest sites (Wilcox et al. 1999) (This
AGFD information is based on a limited study area, a small sample size,
and methods used to describe microhabitat characteristics and may have
only limited applicability in project evaluation); (4) vegetation
providing mid-story and canopy level cover (this is provided primarily
by trees greater than 2 m (6 ft) in height) in a configuration and
density compatible with pygmy-owl flight and dispersal behaviors.
Within 15-m radius plots centered on nests and perch sites, AGFD has
documented the mean number of trees and average height of trees for
Sonoran desertscrub and semidesert grassland areas. The mean number of
trees per plot in Sonoran desertscrub plots was 12.5 with a mean height
of 3.95 m. The mean number of trees in semidesert grassland was 28.5
with a mean height of 8.1 m (Wilcox et al. 2000) (This AGFD information
is based on a small sample size using a method designed to describe
microhabitat characteristics. These numbers may have only limited
applicability in project evaluations); and (5) habitat elements
configured and human activity levels minimized so that unimpeded use,
based on pygmy-owl behavioral patterns (typical flight distances,
activity level tolerance, etc.), can occur during dispersal and within
home ranges (the total area used on an annual basis).
We determined that these proposed primary constituent elements of
critical
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habitat provide for the physiological, behavioral, and ecological
requirements of the pygmy-owl. The first primary constituent element
provides the general biotic communities which are known to support
pygmy-owl habitat in Arizona. We conclude that this element is
essential to the conservation of the pygmy-owl because the species is
not known to occur outside of these biotic communities.
The second primary constituent element provides the components
necessary for nesting, such as cavity availability and cover. The third
primary constituent element describes the structural makeup of habitat
necessary to meet the biological needs of the pygmy-owl such as
breeding, nesting, roosting, perching, foraging, predator avoidance,
and thermal cover, and also promotes prey diversity and availability.
The fourth primary constituent element describes the structural
makeup of vegetation necessary to meet the biological needs of the
pygmy-owl related to movements and dispersal. This includes small-scale
movements for foraging, defense, predator avoidance, pair formation,
nest site selection, etc., as well as landscape level movements needed
to promote genetic diversity and expansion of the population.
The fifth constituent element describes landscape conditions which
may affect pygmy-owl behavioral patterns and relates to the need to
protect habitats from various disturbances. Pygmy-owl behavior is not
typically affected by low levels of human activity or activities which
are predictable (Abbate et al. 1999, 2000, AGFD unpubl. data). Low-
density (< 3 houses per acre) residential areas and roads with low
traffic volumes are examples of this type of activity. However, high
levels of human activities, high-intensity activities, or activities
which cannot be predicted may affect the areas pygmy-owls will use for
nesting, foraging and dispersal (AGFD unpubl. data). High-density
( 3 houses per acre) residential, commercial areas with
lights and constant high levels of activity or unpredictable activities
of any level, ball fields, and roads with high traffic volumes are some
examples of activity levels that could potentially affect pygmy-owl
behavior and habitat use. Habitat elements should be configured, and
human activities should be minimized, so dispersal and pygmy-owl
activities within its home range are not impeded.
We did not map critical habitat in sufficient detail to exclude all
developed areas and other lands unlikely to contain primary constituent
elements essential for pygmy-owl conservation. Within the proposed
critical habitat boundaries, only lands containing some or all of the
primary constituent elements (defined above) are proposed as critical
habitat. Existing features and structures within proposed critical
habitat, such as buildings; roads; residential landscaping (e.g., mowed
nonnative ornamental grasses); residential, commercial, and industrial
developments; and lands above 1,200 m (4,000 ft) do not contain some or
all of the primary constituent elements. Therefore, these areas are not
considered critical habitat and are specifically excluded by
definition.
Facilitating the movement of juvenile pygmy-owls to establish
breeding sites, as well as movements among currently known local
populations of pygmy-owls, is important for dispersal and gene flow,
and providing such connectivity is a widely accepted principle of
conservation biology. Thus, portions of CHUs may function primarily to
provide such connectivity within and among CHUs and may contain only
the primary constituent elements required for dispersal, but we
recognize the essential nature of such connectivity to the persistence
of pygmy-owls in Arizona.
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