Pinyon-juniper woodlands in Zion National Park, Utah

Juniperus osteosperma-Pinus monophylla or P. edulis (P-J) woodlands are the most widespread plant community in Zion National Park (ZNP), southwestern Utah. These woodlands dominate nearly half of the park's land area. Our study of this vegetational complex is based on a sample consisting of 115 macroplots (each 0.01 ha in area) objectively distributed across the entire area of ZNP. We recognize 3 subtypes within the P-J complex: Juniperus osteosperma (Utah juniper) alone, juniper with P. monophylla (single-leaf pinyon), and juniper with P. edulis (two-leaf pinyon). The 2 pinyon pines rarely occur together, and thus the foregoing subtypes do not overlap geographically to a significant extent. The first 2 subtypes occur primarily below 1800 m elevation, while the latter is most commonly found above that elevation. Because of the scarcity of sizable expanses (over ～ 10 ha) of well-developed soils in ZNP, the P-J complex occurs primarily on sites where exposed bedrock covers a large portion of the habitat. As a result, over 90% of stands assigned to the P-J complex support less than 50% tree canopy cover (64% have less than 25% tree cover). Shrub cover increases along the woodland successional gradient. Pinyon cover increases faster than juniper cover. Microbiotic soil crust cover is consistently greater in J. osteosperma-P. monophylla woodlands than in J. osteosperma-P. edulis woodlands, but total living cover increases significantly along the successional gradient in both communities. To enhance plant and animal biodiversity, we recommend that pinyon-juniper woodlands of Zion National Park be managed so that late seral stages do not dominate large tracts.     

Comparison of two roadside survey procedures for dwarf mistletoes on the Sawtooth National Forest, Idaho

Two roadside surveys were conducted for dwarf mistletoes parasitizing lodgepole pine and Douglas-fir on the Sawtooth National Forest, Idaho. One survey used variable-radius plots located less than 150 m from roads. The 2nd survey used variable-radius plots established at 200-m intervals along 1600-m transects run perpendicular to the same roads. Estimates of the incidence (percentage of trees infected and percentage of plots infested) and severity (average dwarf mistletoe rating) for both lodgepole pine and Douglas-fir dwarf mistletoes were not significantly different for the 2 survey methods. These findings are further evidence that roadside-plot surveys and transect-plot surveys conducted away from roads provide similar estimates of the incidence of dwarf mistletoes for large forested areas.     

Late quaternary plant zonation and climate in southeastern Utah

Plant macrofossils from packrat middens in two southeastern Utah caves outline development of modern plant zonation from the late Wisconsin. Allen Canyon Cave (2195 m) and Fishmouth Cave (1585 m) are located along a continuous gradient of outcropping Navajo Sandstone that extends from the Abajo Mountains south to the San Juan River. By holding the site constant, changes in the floral composition for a plot of less than one hectare can be observed, even if sporadically, over tens of millennia. At Allen Canyon Cave, Engelmann spruce-alpine fir forest was replaced by the present vegetation consisting of pinyon-juniper woodland on exposed ridgetops and cliffside stands of Douglas fir, ponderosa pine, and aspen. Xerophytic woodland plants such as pinyon, plains prickly pear, and narrowleaf yucca arrived sometime in the middle Holocene between 7200 and 3400 B.P. At Fishmouth Cave, Utah juniper in Holocene middens replaced blue spruce, limber pine, Douglas fir, and dwarf and Rocky Mountain junipers in late Wisconsin samples. Disharmonious associations for the late Wisconsin occur only at the lower site with the xerophytes Mormon tea, plains prickly pear, and narrowleaf yucca growing alongside subalpine conifers. One possible explanation involves the late Wisconsin absence of ponderosa and pinyon pines from the Colorado Plateaus. Released from competition at their lower limits, subalpine conifers were able to expand into lower elevations and mix with xerophytic plants found today in understories of pinyon-juniper and ponderosa pine woodlands. Quantitative climatic estimates are derived for the late Wisconsin by applying vertical gradients for temperature and precipitation to the amount of vegetation depression. The Fishmouth Cave sequence indicates a minimum lowering of 850 m for blue spruce, limber pine, and dwarf juniper. A depression of at least 700 m for Engelmann spruce and alpine fir is suggested for the Allen Canyon locality. Use of conservatively low gradients for stations below 2080 m yields a 3–4 C cooling from present mean annual temperature and 35 to 60 percent more rainfall than today. Steeper gradients associated with more mountainous terrain suggest a 5 C lowering in temperature and up to 120 percent increase over modern precipitation.        

Fungi associated with soils collected beneath and between pinyon and juniper canopies in New Mexico

The soil fungal community beneath pinyon ( Pinus edulis Engelm.) and one-seeded juniper ( Juniperus monosperma [Engelm.] Sar.) tree canopies is described and compared with fungi from adjacent interspace soils dominated by blue grama ( Bouteloua gracilis [H. B. K.] Lag.). Significantly higher organic matter contents and fungal propagule levels were found in soils beneath pinyon and juniper trees than in interspace soils. Soils under pinyon and juniper trees contained similar chemical, physical, and biological properties and, consequently, many groups of fungi in common (64% of the species isolated were common to both). In contrast soil fungi in adjacent interspace soils were vastly different from those collected in soils beneath pinyon and juniper canopies (44% and 48% species in common, respectively). Soil fungi that were isolated more often from pinyon-juniper soils than from interspace soils included Absidia sspp., Beauvaria spp., Gliocladium spp., Mucor spp., Penicillium cyclopium , P. fasciculata , P. frequentans , P. restrictum , Thamnidium spp., and Trichoderma spp. Soil fungi that were isolated more often in interspace soils than in pinyon or juniper soils included Aspergillus alutaceus spp., A. fumigatus , some Fusarium spp., Penicillium luteum , and P. talaromyces .     

Selection of dwarf mistletoe-infected ponderosa pines by Ips species (Coleoptera, Scolytidae) in northern Arizona

Following an Ips bark beetle outbreak in 2002, mortality of ponderosa pine ( Pinus ponderosa Douglas ex C. Lawson) was evaluated in 2 study areas infested with southwestern dwarf mistletoe ( Arceuthobium vaginatum [Willd.] Presl subsp. cryptopodum [Engelm.] Hawksw. & Wiens) in the Coconino and Tonto National Forests, Arizona. A pairwise comparison of dwarf mistletoe ratings for live and dead ponderosa pines was conducted to determine whether dead ponderosa pines had higher dwarf mistletoe ratings than pines that were not attacked. In both study areas, dead ponderosa pines had significantly higher dwarf mistletoe ratings, indicating an association between the severity of dwarf mistletoe infection and susceptibility to attack by Ips spp. We suggest that the probability of ponderosa pine mortality is greater in stands severely infested with southwestern dwarf mistletoe in northern Arizona.     

Effects of dwarf mistletoe on growth and mortality of Douglas-fir in the Southwest

The effects of dwarf mistletoe ( Arceuthobium douglasii ) on growth and mortality of Douglas-fir ( Pseudotsuga menziesii ) were studied on 387 plots in mixed-conifer stands in three national forests in New Mexico and two in Arizona. Analyses of 8,570 trees showed that low infection ratings (dwarf mistletoe classes 1 or 2) had no significant effect on tree growth, but that losses increased markedly as infection severity increased. Average volume growth losses for trees over 10 inches in diameter were: dwarf mistletoe class 3, 10%; class 4, 25%; class 5, 45%; and class 6, 65%. Mortality of Douglas-fir in stands severely infested with dwarf mistletoe was three to four times that of healthy stands. These high losses confirm the need for silvicultural control of Douglas-fir dwarf mistletoe in the Southwest.     

Succession in pinyon-juniper woodlands following wildfire in the Great Basin

Twenty - one areas in pinyon ( Pinus monophylla ) - juniper ( Juniperus osteosperma ) woodlands burned by wildfire from approximately 1 to 60 years prior to sampling and adjacent unburned mature woodland stands were studied in Nevada and California to determine successional patterns and individual species responses to burning and to changing plant communities through time. One year after burning, all late successional woodland species were present in postburn plant communities except tree species. Increases in both cover and occurrence of annual and perennial forbs resulted in their dominance on early successional sites. Shrubs and annual grasses dominated midsuccessional sites, subsequently giving way to tree, shrub, and perennial grass dominance in late succession. North and east slopes generally supported high cover and occurrence of shrubs, perennial grasses, and perennial forbs, and south and west slopes generally supported high cover and occurrence of annual forbs and annual grasses. The ability to group species according to preferential occurrence on various aspects and successional stages can be used to predict plant community composition in time and space in the pinyon - juniper woodlands.         

Late Pleistocene-early Holocene history of coniferous woodlands in the Lucerne Valley region, Mohave Desert, California

The composition of four radiocarbon-dated, late Pleistocene woodrat middens is reported and analyzed. A date of 11,850 ±550 BP records the first reported macrofossil occurrence in this region of late Pleistocene Pinus monophylla-Juniperus osteosperma woodlands. A 7,800 ±350 BP date documents the most recent J. osteosperma woodlands in this presently coniferless desert area, while a date of 12,100 ±400 BP is the oldest record of juniper woodlands among the four middens. Other juniper and creosote bush desert flora radiocarbon dates along with six pollen profiles were obtained. The research suggests that as recently as 7,800 BP this part of the Mohave Desert was subject to a cooler, moister climate than at present, and that the aboriginal food resources of pinyon seeds and juniper berries were probably available to early prehistoric man in this area.     

Human trampling effects on regeneration and age structures of Pinus edulis and Juniperus monosperma

We examined effects of human foot traffic on age structures and densities of seedlings and saplings of Pinus edulis (Colorado pinyon) and Junisperus monosperma (one-seed juniper) in a heavily used urban park, Garden of the Gods, Colorado. Age structures show no stand-destroying disturbances, but they do contain small peaks 85-95 yr ago (minimum age), which have been interpreted as responses to heavy grazing. For Pinus edulis reverse J-shaped age structures indicate a strongly reproducing population, while flat age structures of J. monosperma show low present reproduction. Young trees showed strong preferences for establishing under existing trees and shrubs and not among herbs or on bare soil. Pinus edulis seedling density was reduced by 73% in heavily trampled areas compared to lightly trampled areas. However, there were no differences in density when only the area protected by rocks, shrubs, or trees was considered. This indicates that direct effects such as physical damage and soil erosion kill young trees, and indirect effects, such as lower seed production, do not cause the lower densities in heavily trampled areas. Pinus edulis saplings and J. monosperma seedlings and saplings showed no differences in density across trampling intensities. In heavily trampled areas of Garden of the Gods Park, recent increases in use have apparently reduced Pinus edulis seedling establishment enough that long-term regeneration is threatened. Managers of all pinyon-juniper woodlands must recognize that in areas strongly impacted by foot traffic, and also presumably by similar disturbances such as vehicle traffic, sufficient regeneration likely does not occur to replace trees. The areal extent of severely disturbed areas should be limited, and managers should seek to avoid further degradation of less damaged areas.     

Infection of young Douglas-firs by dwarf mistletoe in the Southwest

Studies in several areas in Arizona and New Mexico show that dwarf mistletoe ( Arceuthobium douglasii ) is rare in young Douglas-firs growing under infected overstories. Less than 5% of the Douglas-firs under 26 years old and less than 6% of those under 1.4m tall were infected in 77 mistletoe-infested stands. Both percent infection and mean dwarf mistletoe rating of young Douglas-firs increased as tree age, height, and stand dwarf mistletoe ratings increased.     

Effects of dwarf mistletoe on spruce in the White Mountains, Arizona

Mortality of spruce in mixed conifer stands moderately to heavily infested with western spruce dwarf mistletoe was two to five times greater than in healthy stands in the White Mountains, Arizona. Ten-year volume growth loss for heavily infected spruce trees ranged from 25% to 40%. Estimates of growth loss for spruce on a stand basis ranged from 10% to 20% in heavily infested stands. Because western spruce dwarf mistletoe is prevalent in the White Mountains and causes increased mortality and reduced growth, its control should be included in management of mixed conifer stands there.     

Nesting phenology and productivity of birds in the White and Inyo Mountains, California, as assessed with nest-boxes

Nest-boxes were monitored from 1988 to 1992 on 5 grids in the pinyon-juniper ( Pinus-Juniperus ) woodlands of the White and Inyo Mountains, California, to determine breeding phenology and productivity of cavity-nesting birds, and characteristics of used and non-used nest-boxes. We found a total of 112 nests of 6 species. Bewicks Wrens ( Thryomanes bewicki i; 64 nests), Mountain Chickadees ( Poecile gambeli ; 18), Ash-throated Flycatchers ( Myiarchus cinerascens ; 12), and Juniper Titmice ( Baeolophus ridgwayi ; 11) were the most common species utilizing the boxes. Nest phenology and numbers of young were similar to values reported elsewhere in the literature for the species, although nesting success for Mountain Chickadees appeared lower in our study. Chickadees were associated with relatively dense, mature vegetation on southern slopes. Juniper Titmice used areas with tall juniper shrubs and generally sparse vegetation. Bewicks Wrens used areas with short trees, sparse vegetation, and many stumps. Nest-boxes were underutilized (< 15% use) by most species relative to their availability and relative to detected abundances of the species. We suggest that nest-boxes can provide valuable breeding biology information and can potentially increase the productivity of rare and cavity-limited species.     

Species diversity and habitat complexity: does vegetation organize vertebrate communites in the Great Basin?

In this study, we have examined the effect of vegetation structure on the three major vertebrate taxa in Great Basin habitats of southwestern Utah. The effect of increasing vegetation heterogeneity, both horizontally and vertically, on the diversities of lizards, rodents, and postbreeding birds was investigated. We found no statistically significant relationship between diversity of all animal taxa and horizontal vegetation heterogeneity, although lizard diversity tended to decrease with increasing heterogeneity and rodent diversity tended to increase. Bird species diversity was positively correlated with vertical habitat heterogeneity. Abundances were highest for rodents in pinyon/juniper habitat and highest for lizards and birds in areas with the highest grass cover. Species richness was highest in sagebrush habitat for rodents but highest for lizards and birds in pinyon/juniper. Evenness values were relatively similar and high for birds and rodents and were relatively high for lizards in all habitats except for pinyon/juniper, which had an evenness value of 0.38. For rodents and lizards, abundance was significantly correlated with the index for horizontal habitat heterogeneity. After logarithmic transformation, abundance of lizards was positively correlated with increasing vegetation complexity. Combined abundance of lizards and rodents was also positively correlated with vegetation complexity. Rodent and lizard abundances, however, were affected by different aspects of the habitat. After logarithmic transformation, lizard abundances increased significantly with increasing grass cover, whereas rodent abundances increased significantly with increasing shrub cover.     

Attributes associated with probability of infestation by the pi?on ips, Ips confusus (Coleoptera: Scolytidae), in pi?on pine, Pinus edulis

We examined attributes of pi?on pine ( Pinus edulis ) associated with the probability of infestation by pi?on ips ( Ips confuses ) in an outbreak in the Coconino National Forest, Arizona. We used data collected from 87 plots, 59 infested and 28 uninfested, and a logistic regression approach to estimate the probability of infestation based on plot and tree-level attributes. Pi?on pine stand density index was a good predictor of the likelihood of infestation by pi?on ips at the plot level, and a cross-validation analysis confirmed that the model correctly classified 82% of the cases. Diameter at root collar and pi?on dwarf mistletoe infestation level were good predictors of individual tree infestation, and a cross-validation analysis indicated that the model correctly classified 72% of the cases. Results suggest that the occurrence of pi?on ips infestations may be related to stress factors associated with increased stocking and pi?on dwarf mistletoe infestations.     

Wildlife use of Douglas-fir dwarf mistletoe witches' brooms in the Southwest

We evaluated wildlife use of witches' brooms associated with infection by Douglas-fir dwarf mistletoe ( Arceuthobium douglasii ) in 6 mixed-conifer study areas in Arizona and 2 areas in New Mexico. We climbed 153 infected Douglas-firs ( Pseudotsuga menziesii ) and examined 706 witches' brooms for evidence of wildlife use. Even though we observed evidence of use by birds, most wildlife use was by small mammals, particularly red squirrels ( Tamiasciurus hudsonicus ). Red squirrels used witches' brooms for nesting, foraging, caching, and as latrines. Witches' brooms classified as Type II or III brooms, located close to the main bole with large platforms, and 5–10 m above the ground were the most frequently used by red squirrels.     

Diseases associated with Juniperus osteosperma and a model for predicting their occurrence with environmental site factors

On 17 Utah juniper ( Juniperus osteosperma [Torrey] L.) sites studied in Utah, Gymnosporangium inconspicuum was the most common rust fungus, followed in frequency and severity by G. nelsoni, G. kernianum, and G. speciosum . The incidence of G. kernianum was correlated with moderate temperatures and greater than average precipitation. True mistletoe, Phoradendron juniperinum Engelm., was present on seven sites. Incidence of foliage diseases of the mold-mildew type was low on sites with low spring and summer temperatures and high on sites with high summer and fall precipitation. Wood rot was common, and incidence seemed to be correlated with low winter temperatures and low soil nitrate but not with annual precipitation. Needle blight, shoot dieback, and needle cast symptoms were common and considered of abiotic origin. Their nonparasitic nature was indicated by lack of association with pathogenic organisms and the positive correlation of their incidence with winter injury and summer drought factors. Needle blight was also positively correlated with high soil salinity but negatively with high soil calcium regardless of salinity. A nonparametric model was developed that accurately predicted the frequency of the mold-mildew type diseases of J. osteosperma based on measured environmental site factors.     

Habitat of three rare species of small mammals in juniper woodlands of southwestern Wyoming

Southwestern Wyoming constitutes the northern limit of the ranges of the cliff chipmunk ( Tamias dorsalis ), pinyon mouse ( Peromyscus truei ), and canyon mouse ( P. crinitus ). In addition to trying to determine their presence in the region, we wanted to identify habitat characteristics commonly used by each of these species. We used Sherman live-traps to sample 14 sites representing 2 distinct habitat types in 1998 and 1999: juniper-rocky slopes and juniper cliffs. Seventeen habitat characteristics were measured at capture locations for each species and compared with randomly located points. Best subsets multiple logistic regression was used to construct models that distinguish between used and available habitat for each species. The cliff chipmunk occurred in both rocky slopes and cliffs. The pinyon mouse was also captured in rocky slopes and cliffs and was most often captured in locations in the interior of the juniper woodland with high tree canopy cover, high forb cover, and low density of rock outcrops. The canyon mouse was captured only in cliffs at sites consisting of high forb cover, high rock cover, and high tree density.     

Winter habitats and foods of Blue Grouse in the Sheeprock Mountains, Utah

Winter habitat use and food habits of Blue Grouse ( Dendragapus obscurus ) were studied in an isolated Utah desert mountain range that contained little typical Douglas-fir ( Pseudotsuga menziesii ) winter habitat. Habitat use was concentrated in the Douglas-fir and pinyon ( Pinus edulis )-juniper ( Juniperus spp. ) habitat. Douglas-fir and pinyon pine were the most consumed foods. Other foods that represented >15% of the composition of an individual fecal sample were limber pine ( Pinus flexilis ), mahogany ( Cercocarpus ledifoliu ), juniper, and an Anteunaria-Cirsium type. The breadth in winter diet indicates that Blue Grouse may successfully occupy other habitats when typical winter habitat is scarce.     

Juniperus occidentalis (western juniper) establishment history on two minimally disturbed research natural areas in central Oregon

While a trend toward western juniper ( Juniperus occidentalis spp. occidentalis ) super-dominance in big sagebrush ( Artemisia tridentata ) communities of the Pacific Northwest since the late 1800s has been well documented, establishment dates of western juniper in less disturbed environments have not. In this paper we document the establishment history of western juniper on 2 minimally disturbed research natural areas that have substantial differences in their physical characteristics. On each site we randomly established twenty 0.05-ha plots to obtain per hectare counts of mature and juvenile western juniper and to obtain a sample of 100 trees closest to the plot center. These trees were then dated using standard dendrochronological techniques. The lower-elevation, more xeric site has an establishment history that suggests it is an emerging western juniper woodland, with the majority of trees sampled establishing since 1978. The higher-elevation site is an older, well-established woodland with a more even temporal distribution of trees. These results suggest that suitable establishment sites may switch from canopy dependence in emerging woodlands to open sites in mature woodlands and that neither domestic livestock grazing nor active fire suppression is a required mechanism for establishment.     

A 4000-year record of woodland vegetation from Wind River Canyon, central Wyoming

Plant macrofossil analyses of 16 radiocarbon-dated woodrat middens spanning the past 4000 years from the Wind River Canyon region in central Wyoming provide information concerning late Holocene development of juniper woodlands. The study sites are currently dominated by Juniperus osteosperma , with J. scopulorum present locally. Woodlands in the region were dominated by J. scopulorum from ca 4000 yr BP until at least 2800 yr BP. Juniperus osteosperma invaded and expanded before 2000 yr BP. This expansion fits a regional pattern of J. osteosperma colonization and expansion in north central Wyoming during a relatively dry period between 2800 and 1000 yr BP. At the time the Wind River Canyon region was colonized by J. osteosperma , the species had populations 50-100 km to both the north and south. Long-distance seed dispersal was required for establishment in the study area. Genetic studies are necessary to identify source populations and regions.