Dwarf mistletoe-pandora moth interaction and its contribution to ponderosa pine mortality in Arizona

The interaction between Southwestern dwarf mistletoe, Arceuthobium vaginatum subspecies cryptopodum , infestation and defoliation by the pandora moth, Coloradia pandora pandora , on the Kaibab Plateau in Arizona was evaluated. Heavy defoliation of ponderosa pine, Pinus ponderosa , in 1979 and 1981 resulted in mortality of individual trees in areas of heavy dwarf mistletoe infestation. Postmortem evaluation of ponderosa pines indicated that dead trees had a significantly higher dwarf mistletoe rating than did nearby paired live trees. Of 25 tree pairs evaluated, only two live trees had higher dwarf mistletoe ratings than the paired dead tree. Mean dwarf mistletoe ratings were: live trees 2.9, dead trees 4.6 (6 class dwarf mistletoe rating system). Implications for management of the pandora moth are discussed.        

Estimates of the incidence of mistletoes in pinyon-juniper woodlands of the Coconino National Forest, Arizona

The relative incidence of mistletoes found in pinyon-juniper woodlands is estimated for the Coconino National Forest, Arizona, using a roadside survey. Approximately 50% of the pinyon-juniper woodlands surveyed were infested with juniper mistletoes ( Phoradendron juniperinum or P. capitellatum ). In contrast, only about 12% of the areas surveyed were infested with pinyon dwarf mistletoe ( Arceuthobium divaricatum ). In these infested woodlands, 70% with juniper mistletoes and 44% with pinyon dwarf mistletoe were lightly infested. Most areas where pinyon dwarf mistletoe populations were observed (90%) also had juniper mistletoe ( P. juniperinum ) present, but many juniper mistletoe populations observed had no pinyon dwarf mistletoe (50%), although pinyon pine was common at those locations. Therefore, pinyon dwarf mistletoe is frequently co-distributed with juniper mistletoe as has been suggested by other investigators, but juniper mistletoe frequently occurs where there is no pinyon dwarf mistletoe present.     

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.     

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.     

Infection of young Douglas-firs and spruces by dwarf mistletoes in the Southwest

Mistletoe infection of Douglas-fir and spruce seedlings increased as the mean dwarf mistletoe rating of the overstory, seedling density, and total age of seedlings increased. Percent of spruce seedlings infected increased more rapidly than for Douglas-fir as overstory dwarf mistletoe ratings increased. However, the intensity of infection as measured by the mean dwarf mistletoe rating of seedlings, increased at about the same rate for spruce and Douglas-fir. Percent infection of seedlings less than 20 years total age was higher for spruce than for Douglas-fir. Douglas-fir and spruce seedling mortality was from two to three times greater in heavily infested stands than in healthy stands.     

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.     

Forest health impacts of bark beetles, dwarf mistletoe, and blister rust in a Lake Tahoe Basin mixed conifer stand

Interactions between forest health variables and mensurational characteristics in an uneven-aged eastern Sierra Nevada mixed conifer stand were examined. The stand was located in the Lake Tahoe Basin on a site featuring a coarsely textured granitic soil and numerous rock outcrops. Its composition was dominated by California white fir ( Abies concolor var. lowiana [Gord.] Lemm.), with Jeffrey pine ( Pinus jeffreyi Grev. & Balf.) and sugar pine ( Pinus lambertiana Dougl.) less prominent and incense-cedar ( Libocedrus decurrens Torr.) and mountain alder ( Alnus tenuifolia Nutt.) the most minor constituents. The majority of saplings and seedlings were white fir. The stand exhibited no evidence that its development had been influenced by fire and, overall, it consisted of numerous small trees accruing little radial growth. Nearly one-quarter of all standing stems pole size or larger were dead, with mortality concentrated in white fir. Forestfloor fuel accumulations were excessive, and coarse debris was especially prominent. A fir engraver beetle ( Scolytus ventralis LeConte) epidemic in white fir contrasted against apparent endemic population levels of the Jeffrey pine ( Dendroctonus jeffreyi Hopkins) and red turpentine ( Dendroctonus valens LeConte) beetles in Jeffrey pine and of the mountain pine beetle ( Dendroctonus ponderosae Hopkins) in sugar pine. The severity of fir engraver attack on white fir was weakly related to overall tree size and to the proportion of composition consisting of this host species, while in Jeffrey pine and sugar pine, bark beetle attacks were strongly correlated with the individual proportions of these 2 hosts. Across all species, basal area explained a substantial proportion of the variation in overall attack severity. We found light infestations of true fir dwarf mistletoe ( Arceuthobium abietinum Engelm. ex Munz f. sp. concoloris ) in white fir and western dwarf mistletoe ( Arceuthobium campylopodum Engelm.) in Jeffrey pine, plus an early stage of infection by the white pine blister rust ( Cronartium ribicola J.C. Fischer) in sugar pine. Collectively, this case study characterized and quantified many of the conditions, symptoms, and causative agents inherent in a decadent mixed conifer stand in the eastern Sierra Nevada.     

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.     

A seed chalcid ( Eurytoma squamosa Bugbee) parasitizes buckbrush ( Ceanothus fendleri Gray) seeds in a ponderosa pine forest of Arizona

Predispersal seed parasitism rates were quantified for buckbrush ( Ceanothus fendleri Gray) in 3 Arizona ponderosa pine ( Pinus ponderosa Laws.) forest units that had been thinned for ecological restoration objectives. The chalcidoid wasp Eurytoma squamosa Bugbee (Eurytomidae) was responsible for 35% of total seed loss in a single year. These findings represent an expansion of the known range and host list for E. squamosa and increase our understanding of factors that may constrain regeneration of C. fendleri in Southwest ponderosa pine forests.     

Southern range extension of the white fir dwarf mistletoe in Arizona

Discovery of a small isolated population of the white fir dwarf mistletoe, Arceuthobium abietinum f. sp. concoloris, in the Chiricahua Mountains of southeastern Arizona extends the southern distribution of this mistletoe 300 miles. The rare occurrence of the mistletoe in the southwest suggests that it once had a more southerly distribution but only survived past climatic changes in a few locations.     

Home range size of Black-backed Woodpeckers in burned forests of southwestern Idaho

We examined home range size of Black-backed Woodpeckers ( Picoides arcticus ) in burned ponderosa pine ( Pinus ponderosa ) / Douglas-fir ( Pseudotsuga menziesii ) forests of southwestern Idaho during 2000 and 2002 (6 and 8 years following fire). Home range size for 4 adult males during the post-fledging period was 115.6–420.9 ha using the 95% fixed-kernel method, and 150.4–766.1 ha using the 100% minimum convex polygon method. Smoothed bootstrap estimates (95%) were 130.0–521.9 ha. Home range sizes were significantly smaller 6 years after fire than 8 years after fire. Each male had from 2 to 8 areas of concentrated use within his home range. We provide recommendations for estimating area requirements of Black-backed Woodpeckers in post-fire ponderosa pine / Douglas-fir forests.     

Survival and sprouting responses of Chihuahua pine after the Rodeo-Chediski fire on the Mogollon Rim, Arizona

Chihuahua pines ( Pinus leiophylla Schiede and Deppe var. chihuahuana Engelmann) were surveyed on 11 study plots on the Mogollon Rim in east central Arizona to compare characteristics of trees that sprouted from the base or root collar after the Rodeo-Chediski fire with those of trees that did not sprout. The differences in trees killed and top-killed by the fire versus those that survived were also assessed. Trees that sprouted were significantly smaller in height and diameter at breast height; they also experienced lower fire intensities than trees that did not sprout. Smaller trees had higher incidences of mortality than larger trees. These results indicate that, even though Chihuahua pine has fire resiliency, sprouting rates after fire are related to size of trees, age of trees, and burn intensity. Since Chihuahua pine is a rare species in the area studied, its ability to recover from and tolerate fire could prove advantageous for sustainability.     

Ponderosa pine conelet and cone mortality in central Arizona

Ponderosa pine conelets in 10 stands on the Coconino and Kaibab national forests were observed periodically from July 1982 until they matured in September 1983. Abortion, ponderosa pine cone beetles ( Conophthorus ponderosae Hopkins), and ponderosa pine coneworms ( Dioryctria sp., probably Auranticella [Grote]) were the significant mortality factors. Cattle, tip moths, and squirrels rarely destroyed conelets or cones.     

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.        

Production, losses, and germination of Ceanothus fendleri seeds in an Arizona ponderosa pine forest

I quantified seed production and ovule losses for Ceanothus fendleri Gray (Fendler ceanothus) plants protected from large ungulate herbivores in a ponderosa pine ( Pinus ponderosa [Laws]) forest of northern Arizona. I also tested seed germination responses to cold stratification and heat treatments in the laboratory. Fruit production on fecund stems ranged from 7.4 to 38.2 fruits per stem, which equated to 22.2-118.2 potential seeds based on 3 ovules per fruit. Stems that produced fruit were significantly large relative to their expected sizes. Predispersal ovule losses ranged from 70.7% to 91.4% across the 2 years studied. A chalcidoid seed parasite ( Eurytoma squamosa Bugbee) consumed 11%-28% of the total number of seeds produced. Postdispersal seed predation varied from 0% to 24% and was significantly affected by forest floor substrate in 1 study year. Cumulative ovule losses were estimated to be 71%-92%. Cold stratification did not significantly affect seed germination, and exposure to 90°C resulted in the highest germination percentage. Both dormant and nondormant seeds suggested a bet-hedging life history strategy. This study provides basic ecological information important for management of ponderosa pine forest and nursery production of C. fendleri .     

Locality, Habitat, and Elevation Records for the Desert Shrew, Notiosorex crawfordi

Two specimens of Notiosorex crawfordi (Coues) were obtained from Rincon Mountains in southeastern Arizona. Elevations were 2,438 and 2,618 m. At the lower-elevation site the habitat was a meadow in a ponderosa pine forest.     

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.     

Fate and characteristics of Picea damaged by Elatobium abietinum (Walker) (Homoptera: Aphididae) in the White Mountains of Arizona

Spruce aphid, Elatobium abietinum (Walker), is a new invasive pest in high-elevation forests of southwestern USA. Plots in the White Mountains of Arizona were evaluated over several years to assess the extent and severity of damage in high-elevation forests and to identify tree and site characteristics associated with defoliation and mortality. Large areas were defoliated in each of 4 recent outbreaks. Impact from a single defoliation episode included an overall mortality of 10.3% to Picea engelmannii Parry, 24%-41% in severely defoliated trees. Defoliation severity was much greater on P. engelmannii than on P. pungens Engelm. and was more severe in the lower canopy layers. Retention of foliage in the upper-crown third of individual trees was a critical factor in tree survivorship. Mortality was associated with defoliation severity and severe infection by Arceuthobium microcarpum (Engelmann) Hawksworth & Wiens. Picea pungens was much more susceptible to A. microcarpum more than was P. engelmannii . The combined effects of high levels of defoliation and mistletoe infection were lethal, resulting in almost 70% mortality. Mortality continued to occur at 3 years after defoliation. This aphid will affect natural disturbance regimes and tree population dynamics in mixed-conifer and spruce-fir forests of the American Southwest.     

Role of post-Pleistocene dispersal in determining the modern distribution of Abert's squirrel

Abert's squirrel is a forest-dwelling mammal, dependent upon ponderosa pine, that now ranges from southern Wyoming to northern Mexico. During the late Pleistocene, ponderosa pine and this squirrel occurred no further north than central Arizona and New Mexico. In consequence, the present range of the squirrel north of the 36thparallel must have been the result of post-Pleistocene (Holocene) dispersal. If such dispersal took place after the fragmentation of the northern montane conifer forest, at least some leakage across barriers of unsuitable (non-ponderosa pine) habitat must have occurred. Dispersal following transplanting "experiments" has shown that such barriers can be crossed; other evidence is provided that suggests this may occur sufficiently often to produce significant changes in distribution within short periods of time. Thus, explanations for the distribution of Abert's squirrel, based only on historical legacy and local extinctions, are found to be insufficient. An alternative explanation is proposed in which post-Pleistocene dispersal also plays an important role.