Pinyon-Juniper Habitat

1. Habitat Description, Status and Importance

The pinyon-juniper habitat type is one of the most widespread habitats in the southwestern United States (Brown 1994, LaRue 1994), extending over large areas of Utah, Colorado, Arizona, Nevada, and New Mexico (Balda and Masters 1980, Tueller and others 1979; Fig. 1). The total acreage estimates range widely (between 43 and 100 million ac) depending on the definition of pinyon-juniper woodland; the latter figure includes juniper-invaded grasslands. In Arizona, there are approximately 5,328,711 ha (13,167,460 ac) of pinyon-juniper habitat (Brown 1994).

Pinyon-Juniper is a cold-adapted evergreen woodland situated above desert or grassland vegetation and below ponderosa pine forests (Pieper 1977); 1500-2300 m (4650 ft-7130 ft) (Brown 1994). The habitat is characterized by varying co-dominance of juniper and pinyon pine. Junipers are often the more abundant of the two dominant species, but pure stands of either species may occur. Often, as elevation and moisture increase, pinyon pines increase, juniper decrease, total tree density increases, and trees become larger in stature (LaRue 1994, Pieper 1977). Typically, Juniper is dominant at lower elevations with pinyon dropping out completely at the lowest elevation of juniper occurrence.

Several species of juniper are dominant or co-dominant, including Rocky Mountain juniper, Utah juniper, one-seed juniper, alligator juniper, and California juniper. The most common pinyon is Rocky Mountain pinyon, while single-leaf pinyon and Mexican pinyon also occur (Consult Brown 1994, LaRue 1994 and Pieper 1977 for distributional information on the individual species). Understory is variable from completely open to quite dense, especially where sagebrush is present. The stature of pinyon-juniper rarely exceeds 12 m (37 ft) in height. Typically, pinyon-juniper exhibits an open woodland arrangement with well-spaced trees. However, depending on site variables, pinyon-juniper may range from an openly-spaced savanna to a closed forest.

Although soils underlying pinyon-juniper vary, they often are shallow, rocky and low in fertility and are derived from a wide range of parent material including: granite, basalt, limestone, sandstone, and shale (Pieper 1977).

As many as 73 species of birds have been reported to use pinyon-juniper habitat (Balda and Masters 1980). Pinyon-Juniper is also important as a seasonal habitat for elk and mule deer. Human uses of pinyon-juniper are for firewood, pinyon nuts, fence posts, charcoal, railroad ties, mining timbers, and livestock forage (Tueller and others 1979). Increasingly, pinyon-juniper is being recognized for its aesthetic, cultural, threatened and endangered species (Hualapai Mexican Vole, cactus species, 4 threatened and endangered plants (Welch’s milkweed, sentry milk-vetch, Navajo sedge, and Jones’ cycladenia), watershed, and recreational values (Gottfried 1994, Tueller and others 1979). The culture and history of many rural and indigenous populations are connected to pinyon-juniper ecosystems (Gottfried and others 1994).

Wide-scale conversion of pinyon-juniper woodlands to grasslands began after World War II. However, due to fire suppression, large areas of former grassland have also been invaded by juniper. Encroaching juniper are usually found at a lower elevation than pinyon. There is no evidence that pinyon-juniper woodlands with mature pinyon trees 100-200 years old were formerly a grassland invaded by trees (Little 1977).

Conversion was accomplished by various methods including: cabling, bulldozing, hand chopping, grubbing, and burning. Dragging a chain between two dozer tractors was frequently the method of choice. Seeding with grass, especially crested wheatgrass, followed. Widespread conversion has decreased primarily because of high costs and low cost-benefit ratio but also to prevent destruction of archaeological sites (Hart Schwartz pers. comm., Lanner 1981). In Arizona, 485,624 ha (1.2 million ac) of pinyon-juniper were treated in this way from 1950-1961 (Gottfried and others 1994, Little 1977). This conversion occurred in two habitat types: grasslands, mostly at lower elevations where juniper had invaded, and pinyon-juniper woodlands. Conversion of natural pinyon-juniper woodlands to grasslands in the Southwest has included destruction of mature pinyon trees on at least a few hundred thousand acres (Little 1977). Seeding to improve forage has generally proved unsuccessful over large areas and is dependent on annual precipitation, amount of limestone in the soil, pretreatment tree cover, and soil nitrate-nitrogen content (Gottfried and others 1994). In one study, an undisturbed pinyon-juniper stand had greater cover of grasses and forbs than a cabled area after 20 years (Gottfried and others 1994).

The impact of pinyon-juniper conversion on native wildlife has been documented (Swenson 1977 and others). Mule deer and elk use was highest on undisturbed pinyon-juniper (Swenson 1977). The natural pinyon-juniper has wider diversity and higher individuals of bird species (with the exceptions of wintering flocking species) than converted areas (Swenson 1977).

Historic grazing practices have also had an effect both adjacent to and within the pinyon-juniper woodland matrix. These practices have reduced the site potential through soil and vegetation degradation. Soil compaction contributes to or causes increased soil erosion, decreased water infiltration, and reduced soil fertility. The loss of a continuous herbaceous cover especially in adjacent grasslands due to overgrazing has produced a situation where stands do not have enough fuel to carry a fire and eliminate young trees. Fire control has contributed also by allowing small trees to successfully out-compete grasses for water, nutrients and light. Grazing and erosion cause drying of surface soils, which favors deep-rooted species rather than grasses (Gottfried and others 1994).

Selective removal of pinyon will most likely have a serious impact on the breeding bird community (Balda and Masters 1980). Both pinyon and juniper play key roles in maintaining the integrity, survival, and propagation of at least some components of the bird community. Both tree species provide different bird requisites at different times of the year (Balda and Masters 1980).

Removal of trees from illegal fuelwood cutting is also likely to have deleterious effects on the bird species that depend on this habitat. New roads are created from this practice causing increased soil erosion, and removal is usually focused on the large juniper trees which provide the primary food source in this system.

2. Species Descriptions, Objectives and Recommendations

Below are detailed descriptions for each priority bird species in pinyon-juniper habitat. A table at the end of the Pinyon-Juniper section highlights species habitat needs in a quick reference format (Table 10).

Gray Flycatcher (Empidonax wrightii)

Associated Species: Other species that may use similar habitat components or respond positively to management for the Gray Flycatcher are: Plumbeous Vireo, Juniper Titmouse, Bewick’s Wren, Pinyon Jay, Western Scrub-Jay, Black-throated Gray Warbler, Ash-throated Flycatcher, Western Bluebird, and Scott’s Oriole.

Distribution: The Gray Flycatcher breeds in western North America from extreme southern British Columbia (Okanagan Valley), southcentral Washington, central and eastern Oregon, south-central Idaho, and southeastern Wyoming south through western and southern Colorado, eastern California, northern and east-central Arizona, and western New Mexico (AOU 1983). In Arizona, it breeds from the Arizona Strip region and the Navajo and Hopi nations south and east to the Bradshaw Mountains and northeastern Graham and central Greenlee Counties (McCarthey and Corman 1996). Its wintering grounds extend from southeastern California and central Arizona south along the Pacific Slope and interior of Mexico to Nayarit, southern Baja California, and Oaxaca (Howell and Webb 1995). In Arizona, it winters locally along the lower Colorado River, near the town of Kirkland, in the lower Verde River drainage south and east to the town of Sasabe, along the San Pedro River Valley, and very locally to the base of the Chiricahua Mountains (Monson and Phillips 1981).

Ecology: In Arizona, spring migration begins in late March, peaks in late April and early May, and continues with stragglers (rarely) to late May. The primary food for Gray Flycatchers are insects, including: butterflies, moths, bees, grasshoppers, and beetles. The scanning perches are on top of shrubs or small trees, and the flycatching airspaces are close to the ground. The flycatcher often will capture insects on the ground or on low plants (Ryser 1985). From late May through July, nests are placed primarily 0.6 - 3.4 m (2 - 11 ft) high in a shrub or crotch of a juniper or pinyon pine (Terres 1980; ABBA unpubl. data). When nesting in juniper woodlands, the nest is largely made of strips of juniper bark and is therefore well camouflaged (ABBA unpubl. data). Estimated density of Gray Flycatchers ranges from 19-29 pairs per 100 ha (247 ac) (T.W. Haislip in Friedmann and others 1977; LaRue 1994). T.W. Haislip (in Friedmann and others 1977) documented moderate Brown-headed Cowbird nest parasitism in local populations in Oregon. In Arizona, fall migration begins in mid-August and continues through mid-October.

Habitat Requirements: Gray Flycatchers breed in semi-arid woodlands and brushy areas that include pinyon pine and/or juniper woodlands, tall sagebrush/greasewood plains, and open ponderosa or Jeffrey pine forests with pinyon and/or juniper understory. Nesting elevations range from approximately 1400-2300 m (4500-7500 ft), very locally to 2750 m (9000 ft) in Arizona (C. LaRue pers. comm.) and 3350 m (11,000 ft) in California (Small 1994). In Arizona, Gray Flycatchers are most common in larger and taller stands of pinyon pine and/or juniper with open understory sometimes interspersed with sagebrush, cliffrose, and barberry (ABBA unpubl. data). They may need some ground cover to support insect populations for foraging. Gray Flycatchers winter in arid scrub, edge or open riparian woodlands, and mesquite bosques usually below 1400 m (4500 ft) in Arizona.

Habitat and/or Population Objectives:

Population Objective

1. Maintain population density of >7 pairs per 40 ha (100 ac) (Masters 1979 1.8-3.6 pairs/40 ha (100 ac), LaRue 1994 7.6-11.5 pairs/40 ha (100 ac)) in Pinyon-Juniper on the Mogollon Rim and the Colorado Plateau.

Habitat Strategy

1. Manage for pinyon-juniper forests with pinyon to juniper ratio of 1:1 or higher and at least a 13% canopy cover (13%-26% canopy cover, LaRue 1994).

IMPLEMENTATION RECOMMENDATIONS AND OPPORTUNITIES

Management Issues with Conservation Recommendations

Breeding habitat loss and modification of Pinyon-Juniper woodlands has occurred through chaining, clearing, and burning of large, mature woodland tracts for livestock and ungulate forage, house and road development, and fuelwood cutting. Overgrazing by elk and livestock reduces groundcover, inhibits regeneration of shrubs, and increases local cowbird populations. Unitt (1987) suggests there may be an increase in cowbird nest parasitism rates of Gray Flycatchers which may become a serious problem in the future. In Arizona, winter habitat loss includes removal of large tracts of pinyon-juniper woodlands for agriculture, grazing, and fuelwood cutting. Possible threats on wintering grounds in Mexico are largely unknown.

Gray Flycatcher management issues are listed in italics. Below each issue are Arizona Partners in Flight Conservation Recommendations.

Habitat Loss

1. Discourage clearing of large mature tracts of woodland habitat.

2. Encourage small-scale openings.

Grazing

1. Manage grazing pressure (for cattle and elk) to maintain shrub component and grass cover.

Commercial Operations

1. Manage for small-scale openings.

2. Seasonal restriction on fuelwood collection (personal and commercial).

3. Restrict cutting of larger pinyon pines and junipers.

Brown-headed Cowbird Parasitism

1. Maintain appropriate levels of livestock grazing in prime nesting habitat especially during nesting season (May through July).

Implementation Opportunities

1. Consider habitat needs in agency plans and projects, including stewardship projects.

EVALUATION OF ASSUMPTIONS: RESEARCH AND MONITORING

Recommended Research

1. Identify cowbird parasitism rates and their effect on productivity.

2. Identify possible threats on wintering grounds.

3. Quantify breeding habitat.

4. Determine effects of fuelwood harvest, fire, and grazing on habitat requirements.

5. Determine current population density in AZ.

Outreach Needs

1. Educate agency and public of the importance of pinyon-juniper habitat to birds as well as its economic and cultural values.

2. Provide information about the habitat requirements of pinyon-juniper birds to woodcutters and agency personnel.

 

Pinyon Jay (Gymnorhinus cyanocephalus)

Associated Species: Other species that may use similar habitat components or respond positively to management for the Pinyon Jay are: Hairy Woodpecker, White-breasted Nuthatch, Northern Flicker, Cassin’s Kingbird, Mountain Chickadee, Clark’s Nutcracker (foraging).

Distribution: Range of the Pinyon Jay is tied primarily to the distribution of pinyon-juniper woodlands of the Southwest and Intermountain regions of the United States. They breed as far north as central Montana and south to Baja California (Balda and Bateman 1971, Ligon 1978, Marzluff and Balda 1992). In Arizona, Pinyon Jays are permanent residents of pinyon-juniper woodlands and lower ponderosa pine forests in the northern and central part of the state (Balda and Bateman 1971), ranging east to Natanes Plateau, west to the Hualapai Indian Reservation, south possibly to Prescott area, and north to Mount Trumbull (Phillips and others 1964). Pinyon Jays are nonmigratory but may exhibit irregular nomadic movements of hundreds of miles outside normal range during fall and winter when pine seed crops are poor (Balda and Bateman 1971, Phillips and others 1964, Westcott 1964).

Ecology: Pinyon Jays are very early nesters, initiating egg-laying as early as February. Typically, they nest in pinyon-juniper woodlands but will also nest in ponderosa pine forests (approx. 2135 m (7000 ft), Balda and Bateman 1971, Marzluff and Balda 1992). Large flocks (up to 250 individuals) nest communally in traditional breeding areas. Courtship begins in November and pairs form in January-February. Pair bonds are long-term and mates interact throughout the year (Balda and Bateman 1971). Highly synchronous flock nest building begins late February to mid-March. Females incubate, but both parents feed nestlings. Older fledglings are fed by parents and helpers. Young attain independence at 16 weeks. Pairs will renest up to five times in a breeding season if earlier nesting attempts fail (Marzluff and Balda 1992). Most birds breed at age two and have an average lifespan of five years (Marzluff and Balda 1992).

Breeding is apparently triggered by abundant pinyon pine seeds which are harvested in fall and early winter and cached in breeding areas for use during late winter and early spring. Pinyon pine seeds provide the primary source of reproductive energy for nesting Pinyon Jays (Balda and Bateman 1971, Marzluff and Balda 1992). In years following poor pinyon production, breeding is delayed until April or May when other foods, primarily insects, become common (Ligon 1971). Pinyon Jays will also feed on ponderosa pine seed, fruits, eggs, nestlings, lizards. They feed on the ground, in foliage and hawk for insects (Balda and Bateman 1971).

The Pinyon Jay is a gregarious and highly socialized species. Large, highly integrated flocks are maintained year-round and use well-defined home ranges during most years. During poor seed crop years, individuals and flocks have been observed in southern Arizona as well as at treeline in northern Arizona harvesting limber pine seed (Phillips and others 1964, Westcott 1964, Balda and Bateman 1971). Largest flocks (100s to over 1000 birds) (Bent 1964) seen in late summer and winter.

Habitat Requirements: Food availability seems to be the most important factor determining colony breeding site selection (Gabaldon 1979). Open cup nests (usually one nest/tree) are placed in ponderosa pine, pinyon pine, Gambel’s oak, juniper, and occasionally blue spruce trees. Nests are typically 1-8 m (3-26 ft) high and tend to be south-facing (Gabaldon 1979, Marzluff and Balda 1992). Gabaldon (1979) found nest trees were taller and had higher foliage density than surrounding trees. Gabaldon (1979) also found jays avoided trees with abundant pine cones, perhaps because these might attract predators. Many nests were located along roads and Gabaldon (1979) found these nests to have higher reproductive success. Balda and Bateman (1971) studied a well defined flock of about 250 birds which maintained a 21 km2 (8 mi2) home range which included ponderosa pine forest, pinyon-juniper woodland and grassland. This flock used a traditional nesting area of about 95 ha (230 ac) (Balda and Bateman 1971).

Communal seed caching areas are discrete and located within a flock’s home range. Generally, cache sites are sparsely vegetated, have good drainage and a southern exposure. Thus, these areas are snow-free or first to melt. Birds also tend to cache seeds close to tree trunks where less snow accumulates. Not only do these sites allow for easy retrieval of cached seeds during the early nesting season, but they also provide good conditions for seed germination. Many cached seeds are not consumed and germinate (Ligon 1971). Balda (1987:525) described the relationship between the pinyon jay and pinyon pines as "...one of the best coevolved, mutualistic plant-vertebrate examples known...".

Habitat and/or Population Objectives:

Population Objective

1. Maintain an increasing or stable population trend and distribution throughout pinyon-juniper woodlands in the Colorado Plateau and the Mogollon Rim physiographic areas.

Habitat Strategies

1. Maintain large, cone bearing pinyon trees (75 years or older, Little 1977) in a minimum of 7 sq mi patches (Balda and Bateman 1971) in mature pinyon juniper woodlands or pure pinyon pine woodlands.

 

IMPLEMENTATION RECOMMENDATIONS AND OPPORTUNITIES

Management Issues with Conservation Recommendations

According to Breeding Bird Survey data, Arizona had the highest average statewide density for the Pinyon Jay from 1965-1979 (Robbins and others 1986). However, analyses of these data did not reveal any significant trends for this species. Balda and Marzluff’s (1992) data for an intensively studied pinyon jay population in Flagstaff, from 1972-1986, indicated a declining population.

Three major factors, which vary annually, affect the long-term success of Pinyon Jay populations: size of pinyon pine crops, amount of nest predation, and harshness of the physical environment, particularly the amount of snow during the nesting season (Marzluff and Balda 1992). Although we have no control over the latter, the first two factors can be influenced by human activities. Primary management concerns related to these include: 1) habitat loss due to urbanization, as documented in the Flagstaff vicinity (Marzluff and Balda 1992), as well as to management of pinyon-juniper woodlands (e.g. chaining, burning) and potential habitat loss from Ips beetle invasion of stressed pinyon trees, 2) abundance of mature pinyon pine trees which provide the primary source of food for breeding pinyon jays and which can also be affected by land management practices, and 3) increasing numbers of American Crows and Common Ravens (important nest predators) in Pinyon Jay breeding areas near urban areas (also documented in the Flagstaff area) (Marzluff and Balda 1992).

Pinyon Jay management issues are listed in italics. Below each issue are the Arizona Partners in Flight Conservation Recommendations.

Nest Predation

1. Consider local Common Raven control if their increased numbers are affecting nest success.

Habitat Loss/Habitat Assessment

1. Maintain extensive stands of pinyon with emphasis on cone-producing trees.

2. Limit collection of cone-producing pinyon trees for fuelwood (75 yr or older, Little 1977).

3. Identify and retain traditional home ranges.

4. Inventory pinyon-juniper structural stage distribution to determine how many mature stands (preferred by Pinyon Jay) exist.

5. Reduce Ips beetles by reducing the number of slash piles (winter hibernaculums).

Soil Erosion

1. Encourage small-scale openings to reduce erosion in denser, mature stands.

2. Use appropriate livestock and/or wild ungulate stocking rates or densities to promote grass and herbaceous growth.

Implementation Opportunities

1. Consider habitat needs in Agency plans and projects.

EVALUATION OF ASSUMPTIONS: RESEARCH AND MONITORING:

Recommended Research

1. Determine amount of mature pinyon woodlands in Arizona.

2. Determine if Common Raven nest predation is a serious problem.

3. Determine the effects of fragmentation of nesting stands on Pinyon Jays.

4. Determine the landscape ecology configuration needs for species (i.e. fragmentation, edge effects).

5. Evaluate the effects of human pine nut harvest on Pinyon Jay’s food availability.

Outreach Needs

1. Educate agency and public of the importance of pinyon-juniper habitat to birds as well as the economic and cultural values.

2. Give information about the habitat requirements of pinyon-juniper birds to woodcutters and agency personnel.

3. Educate agency and public on the unique traits of Pinyon Jays (i.e. communalism/mutalism, the "Johnny Appleseed" of the bird world).

 

Gray Vireo (Vireo vicinior)

Associated Species: Other species that may use similar habitat components or respond positively to management for the Gray Vireo are: Ash-throated Flycatcher, Juniper Titmouse, Bushtit, Bewick’s Wren, Blue-gray Gnatcatcher, Black-chinned Sparrow, and Scott’s Oriole.

Distribution: The Gray Vireo breeds from Southern California (locally) and northwestern Baja California, southern Nevada, southern Utah, and southern Colorado south through western and central New Mexico, and isolated localities in the Texas and Oklahoma panhandles, and also southwestern Texas and northwestern Coahuila (A.O.U. 1983, Small 1994, Andrews and Richter 1992, Wauer 1973, Phillips and others 1964). In Arizona, Gray Vireos breed at mid-elevations from the Grand Canyon region east across the Navajo Nation and south through the Mogollon escarpment into the southeastern portions of the state (ABBA unpubl.data, Brown and others 1984, LaRue in prep, Phillips and others 1964). Gray Vireos are rare migrants throughout the state (Brown and others 1984, Phillips and others 1964). They winter mainly in northern Mexico, southern Baja California and rarely in southern Arizona and the Big Bend region of Texas (AOU 1983, Howell and Webb 1995, Wauer 1973).

Ecology: The Gray Vireo arrives in southern Arizona in early April and northern Arizona in late April. They depart these regions in early and late September respectively (LaRue in prep, Phillips and others 1964). Gray Vireos are primarily insectivorous during the breeding season. During the winter, they are frugivorous and rely almost entirely on fruit of elephant trees (Bates 1992). They typically nest low in a small tree or shrub 0.5-2.0 m (2-6 ft) above ground (Ehrlich and others 1988). Young fledge at 13-14 days. Gray Vireos are known hosts of the Brown-headed Cowbird. Gray Vireos tend to occur at naturally low population densities.

Habitat Requirements: Gray Vireos breed in Arizona in open mature pinyon-juniper woodlands on canyon and mesa slopes from 975-2075 m (3200-6800 ft) in elevation. A broadleaf shrub component is typically present, often comprised of Utah serviceberry and single-leaf ash. Gray Vireos may also breed in situations dominated by a chaparral component (T. Corman, AGFD, pers. observ.). In northeastern Arizona, they were absent from woodland stands greater than 280 trees/ha (2.5 ac) (LaRue 1994).

Habitat and/or Population Objectives:

Population Objectives

1. Maintain stable or increasing populations across their range in Arizona.

Habitat strategy

1. Maintain an open pinyon-juniper woodland with a shrubby understory, especially on moderate rocky slopes.

IMPLEMENTATION RECOMMENDATIONS AND OPPORTUNITIES

Management Issues with Conservation Recommendations

Apparently some population declines of Gray Vireos have been noted in California (Small 1994) and the species is on Arizona’s Wildlife of Special Concern list (AGFD 1996 draft). Although it is a known cowbird host, no negative impacts have been clearly identified at this time. The apparent extreme winter dietary specialization as well as the tendency to occur in low densities, confers some intrinsic vulnerability which could result in population declines. Because of their tendency to occupy undisturbed canyon and mesa slopes, Gray Vireos may be relatively immune to habitat-related population declines. In general, life history of the Gray Vireo is still poorly known.

Gray Vireo management issues are listed in italics. Below each issue are the Arizona Partners in Flight Conservation Recommendations.

Fire Suppression

1. Manage fire to maintain existing gray vireo habitat matrix and to prevent stands from becoming too dense.

Brown-headed Cowbird Parasitism

1. Discourage development of additional livestock water sources to reduce the number of cowbirds in Gray Vireo habitat.

2. Discourage highly intensive, short-term grazing that may greatly alter habitat structure and increase the presence of cowbirds.

Implementation Opportunities

1. Consider habitat management needs of Gray Vireo in agency plans and local projects.

EVALUATION OF ASSUMPTIONS: RESEARCH AND MONITORING

Recommended Research

1. Determine the effects of Brown-Headed Cowbird parasitism.

2. Study Gray Vireo habitat selection.

3. Collect missing general natural history of Gray Vireo.

4. Determine natural fire regime/interval in successful breeding areas.

5. Restart the Forest Service pinyon-juniper initiative to acknowledge the benefits and uses of pinyon-juniper habitat.

Outreach Needs

1. Educate agency and public of the importance of pinyon-juniper habitat to birds as well as the economic and cultural values.

2. Give information about the habitat requirements of pinyon-juniper birds to woodcutters and agency personnel.

 

Black-throated Gray Warbler (Dendroica nigrescens)

Associated Species: Other species that may use similar habitat components or respond positively to management for the Black-throated Gray Warbler are: Plumbeous Vireo, Juniper Titmouse, Bewick’s Wren, Pinyon Jay, Western Scrub-Jay, Ash-throated Flycatcher, Western Bluebird, and Scott’s Oriole.

Distribution: The Black-throated Gray Warbler’s breeding range extends from southwestern British Columbia south through the coastal states to northern Baja California. Eastward it extends from eastern Oregon through southern Idaho and Wyoming south to southeastern Arizona, southwestern New Mexico, extreme west Texas (breeding status unknown in Guadalupe Mountains), and northeastern Sonora, Mexico (Dunn and Garrett 1997, Guzy and Lowther 1997). The Black-throated Gray Warbler winters in Baja California, on the Pacific Slope and interior of Mexico, and in small numbers along the West and Gulf coasts of the United States (Guzy and Lowther 1997). In Arizona, this species is found breeding north of the Mogollon Rim and south through eastern Arizona, west to the Baboquivari and Bradshaw Mountains, Grand Canyon Region, and the Hualapai Mountains (ABBA unpubl. data, Monson and Phillips 1981). The Black-throated Gray Warbler is an uncommon winter resident in Phoenix, Tucson, and the Babaquivari Mountains (Monson and Phillips 1981).

Two races of Black-throated Gray Warblers are distinguished by differences in wing length, amount of white in tail, and song. Dendroica nigrescens nigrescens breeds from northwestern California to southwestern British Columbia, and D. n. halseii breeds in eastern Oregon and Washington south through Arizona, New Mexico, and Sonora, Mexico (Morrison 1990, Oberholser 1930). The former race winters in California and Arizona south to northern Mexico, while the latter winters only in Mexico (Morrison 1990). Some authors recognize these two races as distinct subspecies: genetic differences between Black-throated Gray Warblers of Washington and Arizona are as great as those between Townsend’s warblers and hermit warblers (Bermingham and others 1992).

Ecology: The Black-throated Gray Warbler is a short-to-medium-distance Neotropical migrant whose migration route follows the coast and mountain ranges of western North America (Curson and others 1994, Guzy and Lowther 1997). Spring arrival dates in southern Arizona range from mid-March through May, and departure dates range from late July through October. Spring arrival dates in northern Arizona range from mid-Aprilthrough May, and departure dates range from mid-July through early October (Phillips and others 1964). Nesting records from Arizona include: 12 nests with eggs found 4 May-19 June, with the majority of these nests found between 17 and 26 May (n = 7; Bent 1953); an occupied nest on 15 May 1993 in the Hualapai Mountains; a nest with young on 28 May 1995 in Coconino National Forest; and a nest with young on 10 July 1997 north of the Kaibab National Forest on the Arizona (ABBA unpubl. data). Breeding Bird Atlas data suggest that the Black-throated Gray Warbler’s breeding season begins in April in the southern part of their range (an adult was seen carrying food on 9 May 1993, and an adult was seen feeding recently fledged young on 19 May 1995) and extends into August (recently fledged young observed on 7 August 1996). The Black-throated Gray Warbler builds a deep cup nest of leaves, cocoons, oak mast, paper shreds, bark, and other plant material, and it is frequently lined with small feathers of other bird species (Harrison 1979). They typically raise one brood a year, though they may double-brood in some areas (e.g. Monterey County, California; Roberson and Tenney 1993).

The Black-throated Gray Warbler’s diet consists almost exclusively of insects, especially caterpillars (Dunn and Garrett 1997). They primarily forage at the mid-canopy level by gleaning foliage, or occasionally by hover gleaning and sallying for flying insects (Dunn and Garrett 1997). This species is not social during the breeding season, but will join mixed-species flocks with other insectivorous birds during winter and migration (Dunn and Garrett 1997). Known predators of adults include Sharp-shinned Hawks and Cooper’s Hawks (Reynolds and Meslow 1984), and likely predators of eggs and young include jays, crows, and snakes (Bent 1953, Grinnell and Storer 1924). There is little information on the extent which brown-headed cowbirds affect Black-throated Gray Warblers. Bent (1953) reported that brood parasitism was not a problem for this species, but recent reports suggest that parasitism rates are higher than previously thought or are increasing. Research from four different locales in the western United States suggests parasitism rates between 11% and 21% (see Guzy and Lowther 1997). Thirteen percent of Black-throated Gray Warbler family groups (n = 30) reported by the Arizona Breeding Bird Atlas (ABBA unpubl. data) had a fledgling cowbird.

Habitat Requirements: In northern Arizona, the Black-throated Gray Warbler is primarily associated with pinyon pine and juniper woodlands (occasionally with scattered ponderosa pine) and mixed oak-pine woodlands. In southern Arizona, this species occupies oak-alligator juniper woodland, Chihuahuan pine, Mexican pinyon pine, Emory oak, and Arizona white oak, as well as other mixed oak-conifer associations along canyons and steep slopes (Balda 1969, Dunn and Garrett 1997). Breeding habitat is frequently characterized by a brushy undergrowth of scrub oak, ceanothus, manzanita, or mountain mohagany (Dunn and Garrett 1997). During spring and fall migration, these warblers can be found in a variety of forest, woodland, scrub, and thickets similar to that used during the breeding season (AOU 1983), as well as desert washes and desert riparian areas (Troy Corman, pers. observ.). Individuals that winter in Arizona are primarily associated with cottonwood-willow and sycamore-mesquite vegetation (Monson and Phillips 1981). In addition, Black-throated Gray Warblers have become more common as winter residents in shade trees of urban areas, such as Phoenix and Tucson (Troy Corman, AGFD, pers. observ.).

Little information is available on microhabitat characteristics of nest sites. Nests are typically placed on a horizontal tree branch or near the main stem of a shrub (Harrison 1979). Of seven nests found in southeastern Arizona, six were in white or Emory oak and one was in juniper, average nest height was 7.5m (24.5 ft) (range 3.6-12.2 m or 12-40 ft), and nests were 1.2-3.0 m (4-10 ft) from the tree trunk (Harrison 1984). Other nests found in Arizona include one from the Chiricahua Mountains that was in a dense mistletoe clump, 0.46 m (1.5 ft) high and 0.86 m (3 ft) from the trunk of a scrub oak, and one from northern Arizona that was 3.25 m (10.5 ft) high in a juniper tree and 0.78 m (2.5 ft)from the trunk (ABBA unpubl. data).

Habitat and/or Population Objectives:

Population Objective

1. Maintain a population density of 11.5 pairs /40 ha (100 ac) (7.6-15.3 pairs/40 ha, LaRue 1994) in Pinyon-Juniper woodlands on the Mogollon Rim and Colorado Plateau.

Habitat Strategy

1. Manage for pinyon-juniper forests with a pinyon to juniper ratio of 1:1 or higher and at least a 13% canopy cover (13%-26% canopy cover, LaRue 1994).

IMPLEMENTATION RECOMMENDATIONS AND OPPORTUNITIES

Management Issues with Conservation Recommendations

There is little information on overall population trends for the Black-throated Gray Warbler. Breeding Bird Survey (BBS) data suggest steady or slightly increasing numbers from 1966-1991 (Peterjohn and others 1995). This species does not appear to be greatly impacted by human activities and will occupy areas that have been altered. However, there have been no detailed studies of responses to habitat alteration, such as changes in densities, breeding success, and habitat use (Guzy and Lowther 1997). Techniques used for improving pasturelands, such as the removal of overstory trees from pinyon-juniper woodland, may adversely affect habitat use by Black-throated Gray Warblers (Sedgwick 1987). Continued alteration and loss of habitat may have cumulative effects unidentified to date. For example, land management practices that increase contact between Black-throated Gray Warblers and brown-headed cowbirds may have a substantial impact on breeding success.

Black-throated Gray Warbler management issues are listed in italics. Below each issue are the Arizona Partners in Flight Conservation Recommendations.

Habitat Loss

1. Discourage clearing of large mature tracts of habitat.

2. Encourage small-scale openings in pinyon juniper woodlands.

Brown-headed Cowbird Parasitism

1. Manage livestock numbers to reduce the number of cowbirds in pinyon-juniper woodlands.

2. Discourage highly intensive, short-term grazing that may alter habitat structure and increase the presences of cowbirds.

Commercial Operations

1. Limit seasonal cutting of pinyon trees (May through July), especially larger sized trees.

Ips Beetles Outbreak

1. Reduce Ips beetles by reducing the number of slash piles (winter hibernaculums).

Implementation Opportunities

1. Educate agency and public of the importance of pinyon-juniper habitat to birds as well as the economic and cultural values.

2. Give information about the habitat requirements of pinyon-juniper birds to woodcutters and agency personnel.

3. Consider Black-throated Gray Warbler habitat needs in Agency plans and projects.

EVALUATION OF ASSUMPTIONS: RESEARCH AND MONITORING

Recommended Research

1. Research the general natural history of the Black-throated Gray Warbler including: breeding biology, foraging biology (species role in limiting number of devastating insects in pinyon), and habitat requirements.

2. Determine habitat selection parameters to assess how fuelwood harvest may affect the Black-throated Gray Warbler.

3. Determine cowbird parasitism rates and effects on Black-throated Gray Warblers.

Juniper Titmouse (Baeolophus griseus)

Associated Species: Other species that may use similar habitat components or respond positively to management for the Juniper Titmouse are: Ash-throated Flycatcher, Gray Vireo, Pinyon Jay, Western Scrub Jay, Black-throated Gray Warbler, Western Bluebird, Scott’s Oriole.

Distribution: Resident from southeastern Oregon, northeastern Nevada, southeastern Idaho, southern Wyoming, central Colorado, and extreme Oklahoma south (east of the Sierra Nevada) to southeastern California, central and southeastern Arizona, extreme northeastern Sonora, southern New Mexico, and extreme western Texas (AOU 1998). In Arizona, it is a fairly common to common resident in the northeastern, northern, central, and locally southeastern portions of the state. The range extends west to Mount Trumbull and the Cerbat, Hualapi, Bradshaw, Superstition, Galiuro, and Chiricahua Mountains (Monson and Phillips 1981).

Ecology: An obligate inhabitant of pinyon-juniper woodlands (Andrews and Righter 1992, Behle 1985, Phillips and others 1964, Small 1994). Occurs as singles or pairs and does not typically form conspecific flocks although it does occur in mixed-species flocks (Phillips and others 1964). Balda (1987) states that Juniper Titmouse are "major pine seed predators" that may consume "large numbers of seeds." Bradfield (1974) observed it feeding on juniper seeds in the fall. It is likely largely insectivorous during the warmer half of the year. An obligate secondary cavity nester. Of 13 active nests found as part of the Arizona Breeding Bird Atlas, nine (79 %) were in junipers (T. Corman, AGFD, pers. observ.). Nesting dates ranged from 15 May to 30 June. Nest cavity heights were from 1.12 m to 4.40 m. The diameter (dbh) of the nest trees varied from 14-48 cm (5.5-1.5 in). It is probably not subject to brood parasitism by Brown-headed Cowbirds. Breeding densities from three study sites over two years in central Arizona ranged from 28.7 to 52.0 pairs per 40 ha (100 ac) which made up 23.5% to 43.6% of the total breeding bird density (Masters 1979). In a similarly study using identical methods in northeastern Arizona (LaRue 1994) reported 7.6 to 11.5 pairs per 40 hectares comprising 7.4% to 17.7% of the total breeding bird density.

Habitat Requirements: The Juniper Titmouse is highly restricted to pinyon-juniper woodlands (Andrews and Righter 1992, Balda and Masters 1980, Behle 1985, Bradfield 1974, Phillips and others 1964, Small 1994). It occasionally wanders into other habitats (usually riparian) within its range that are adjacent to or near pinyon-juniper woodlands during the nonbreeding season (Andrews and Righter 1992, Bradfield 1974, Brown and others 1984, Phillips and others 1964, Small 1994, Sogge and others 1998). The Juniper Titmouse is virtually unknown as a transient outside of the range cited above (Rea 1983, Rosenberg and others 1991, Witzeman and others 1997). Tree density in two Pinyon-juniper breeding bird investigations that examined stands supporting breeding titmice (LaRue 1994, Masters 1980) ranged from 155 to 380 trees per hectare. Canopy cover of one study (LaRue 1994) varied from 11% to 26%. Combined, these studies indicate that the proportion of the breeding bird density the titmouse contributes to tends to drop with increasing tree density, increasing total bird density, increasing proportion of junipers, and increasing canopy cover.

Habitat and/or Population Objectives:

Population Objective

1. Maintain a stable or increasing population trend within current range and distribution.

IMPLEMENTATION RECOMMENDATIONS AND OPPORTUNITIES

Management Issues with Conservation Recommendations

Formerly known as Plain Titmouse (Parus inornatus). However it has recently been split (AOU 1997) into two species, with the interior forms being called the Juniper Titmouse and those populations west of the Sierra Nevada called the Oak Titmouse (B. inornatus). Most available information on the "Plain Titmouse" (e.g. Ehrlich and others 1988) is based on studies of the Oak Titmouse in California. Therefore, little is known specifically for the Juniper Titmouse. Because it is clearly associated with mature pinyon-juniper woodlands, management activities that favor these stands will benefit this species. Investigations to determine specific habitat requirements and basic natural history are needed.

Juniper Titmouse management issues are listed in italics. Below each issue are the Arizona Partners in Flight Conservation Recommendations.

Habitat Loss

1. Discourage clearing of large mature tracts of habitat.

2. Encourage small-scale opening of habitat.

EVALUATION OF ASSUMPTIONS: RESEARCH AND MONITORING

Recommended Research

1. Determine specific habitat requirements, habitat use and basic natural history for this subspecies.

 

3. Coordination of Recommendations and Opportunities in Pinyon-Juniper

The key issues for pinyon-juniper birds seem to stem from habitat changes over the past 50 years. This conversion has resulted in increased livestock grazing, and consequently, a reduction in ground cover and shrub regeneration, and an increased presence of Brown-headed Cowbirds. Three of the priority species in pinyon-juniper habitat are cowbird hosts and parasitism rates appear to be increasing for all three species (Gray Flycatcher, Gray Vireo and Black-throated Gray Warbler). Although there is no evidence to date that these species are declining from parasitism, it is suggested that adverse effects are likely if the rate of parasitism continues to increase. Loss and/or alteration of habitat, especially larger cone-bearing pinyons, is the primary concern for the fourth species, the Pinyon Jay.

Clearing large tracts of mature trees using chaining, bulldozing, or cabling methods are not common management practices anymore. However, they do still occur in some areas. Loss of habitat today is more likely to be caused by lack of fire, fuelwood cutting of larger trees, and from overgrazing that prevents shrub regeneration. All four priority species suffer from loss and alteration of pinyon-juniper woodlands. It is recommended that seasonal restrictions on fuelwood collections be implemented for both personal and commercial use, and that limits on collection of larger trees, especially cone-producing pinyon (>75 yrs), also be set.

Whether to burn is a question that is being asked across the Southwest over many habitats. In pinyon-juniper, fire suppression has resulted in stands becoming extremely dense causing a reduction in the herbaceous and shrub layer and an increase in soil erosion. All four priority species use the shrubby component in pinyon-juniper habitat and prefer openings between older, taller trees. Burning or mechanical thinning that creates small openings but retains the larger trees is recommended for this habitat. Openings can alleviate soil erosion by allowing the herbaceous layer to grow and stabilize the shallow rocky soils common to pinyon-juniper woodlands. Burning of slash piles is also recommended to eliminate winter hibernaculums of the Ips beetle, that commonly target pinyon trees that are stressed due to drought or over crowding by junipers.

Urbanization has had the most negative effect on the Pinyon Jay. More and more developments moving into existing Pinyon Jay breeding areas have eliminated important habitat and mature cone producing trees that are essential to Pinyon Jay survival. Urbanization has also brought an increased number of crows and ravens, the primary predators of the Pinyon Jay. Inventory of the existing stands of mature pinyon-juniper is recommended to better assess limitations to pinyon tree harvest, if necessary.

 

Table 10. Pinyon-Juniper Priority Species Habitat Needs

Priority Species

Vegetation Composition

Vegetation Structure

Abiotic Factors

Landscape Factors

Gray Flycatcher -primary: pinyon pine and/or juniper, with an open overstory of ponderosa

-secondary: sagebrush, greasewood

-larger stands of PJ with open understory, some areas with sagebrush,

-nest height 0.5-3.0 m (2-9 ft)

-may need some ground cover to support insect populations for foraging

-larger taller stands of sagebrush and greasewood

-elevation 1375-2285 m (4500-7500 ft), locally to 2750 m (9000 ft) -mid to late successional stages

-edge effect and fragmentation do not appear to be an issue

Pinyon Jay -breeds in pinyon and ponderosa pine

-usually in pinyon-juniper where pinyon is dominant

-over 85% of nests found in bottom half of canopy (Balda and Bateman 1971)

-commonly in extensive stands of pinyon-juniper with open physiognomy

-may increase as mid and understory decrease

-nest and cache on south side of trees

-elevation 1525-2285 m (5000-7500 ft)

-may key in onwarmest microclimate for nesting

-mid-late successional (pine nuts in mature trees)

-use extensive stands for foraging, colony may have up to a 13 sq km (8 sq mi) home range (Balda and Bateman 1971)

Gray Vireo pinyon-juniper with broad-leafed shrubs - Utah serviceberry, single-leaf ash -open, not in stands greater than 280 trees/ha (2.5 ac)

-usually nest and forage at <2 m (29 in.-8 ft) (CA FS)

-rocky, drier sites

-moderate to steep slopes (canyon/mesa slopes),

-elevation 975-2075 m (3200-6800 ft)

-not usually found in chained/young pj;

-patch size small.

-Plumbeous Vireo move in when structure is denser, patch size larger.

-need more info.

Black-throated Gray Warbler

 

-mostly pinyon

-also commonly occurs in Madrean oak/ pine-oak in southeastern AZ w/ shrub component

-in taller and denser PJ woodland

-usually nest 2'-15' (0.6-4.5m)(Zeiner and others 1990)

-low to mid-story nester.

-prefers relatively heavy conifer cover (Morrison 1982)

-forage most often in pinyon (LaRue pers. comm.)

-not found where juniper becomes dominant.

-in PJ, usually between 1980-2440 m (6500-8000 ft) in AZ.

-Locally below 1980 m (6500 ft) in PJ.

-commonly found in lower elevations in se AZ habitats.

-may prefer woodlands w/ interspersed shrubby openings

-successional stage: mid to late pinyon woodland

-unknown if fragmentation has an effect on species.

Juniper Titmouse -pinyon-juniper woodlands

-may use riparian habitat if adjacent to pinyon-juniper

-taller pinyon and juniper trees. -drop out with increasing tree density or too few trees late successional pinyon-juniper woodlands

 

Table 11. Special Factors for Pinyon-Juniper Priority Species

Priority Species

Special Factors

Gray Flycatcher -Brown-headed Cowbird host (maybe increasing)

-insectivore low forager - often ground gleaner

-possibly semicolonial

-poorly represented by Breeding Bird Survey (BBS) routes

-a high priority species for most states it breeds in

Pinyon Jay -roost and nest colonially up to 250 individuals

-only one nest per tree, usually

-communal feeders of fledglings between 3-6 weeks old

-long-term pair bonds

-co-evolved with pinyon trees

-may suffer from common raven predation

Gray Vireo -frequent cowbird parasitism

-low foliage gleaner for insects

Black-throated Gray Warbler -Brown-Headed Cowbird parasitism occurs, but effect unknown

-forages low to mid canopy, foliage gleaner

Juniper Titmouse -pinyon-juniper obligate

-occurs mainly as single or pairs but not flocks

-consume large quantities of pine seeds

-secondary cavity nester