Composition of forest stands used by White-headed Woodpeckers for nesting in Washington

In this study, I examined the composition of managed ponderosa pine ( Pinus ponderosa ) forests used by nesting White-headed Woodpeckers ( Picoides albolarvatus ) along the eastern slope of the Cascade Range in Washington. I sampled trees and snags using the point-centered quarter method to assess species composition, tree and snag density, and stand basal area in 16 forest stands containing White-headed Woodpecker nests. All stands had a history of timber management and 2 had been burned and salvage-logged. Mean live-tree density (≥10.16 cm dbh) was 182.3 trees · ha –1 (SE = 13.52), mean snag density (≥10.16 cm dbh) was 11.5 snags · ha –1 (SE = 1.92), and mean stand basal area was 17.2 m 2 · ha –1 (SE = 1.58). Ponderosa pine had the highest importance value (mean = 220.9, SE = 17.25) of any tree species in all but 2 stands. Mean dbh of ponderosa pines was 33.0 cm (SE = 0.26) and ranged from 26.1 to 50.2 cm within stands. Mean density of ponderosa pine was greatest in the 20.3–30.5 cm dbh size class and lowest in the 50.8–61.0 cm and >61.0 cm dbh size classes. Tree density was up to 5.3 times greater than densities believed to be typical of ponderosa pine forests prior to fire suppression. Snag densities were within the range estimated for historical dry forests of the eastern Cascades, yet only 50% of all snags sampled had a dbh >25.4 cm. Although White-headed Woodpeckers are considered strongly associated with old-growth ponderosa pine, my results suggest that they may be more adaptable to using forests dominated by smaller diameter trees. En este estudio, examiné la composición de los bosques manejados de pino ponderosa ( Pinus ponderosa ), utilizados para anidamiento por el pájaro carpintero cabeciblanco ( Picoides albolarvatus ), a lo largo de la vertiente oriental de la cordillera Cascade del Estado de Washington. Muestreé árboles vivos y muertos usando el método de cuadrantes al punto central en 16 rodales con nidos del pájaro carpintero cabeciblanco para evaluar la composición de especies, la densidad de árboles vivos y muertos y el área basal del rodal. Todas las áreas tenían una historia de manejo maderable y 2 habían sido quemadas y taladas para recuperar madera. La densidad promedio de árboles vivos (≥10.16 cm DAP) fue 182.3 árboles · ha –1 (DE = 13.52), la densidad promedio de árboles muertos (≥10.16 cm DAP) fue 11.5 árboles muertos · ha –1 (DE = 1.92) y el área basal promedio de los rodales fue 17.2 m 2 · ha –1 (DE = 1.58). El pino ponderosa tuvo el valor de importancia más alto (promedio = 220.9, DE = 17.25) de las especies de árboles en todos, menos 2 rodales. El DAP promedio de los pinos ponderosa fue 33.0 cm (DE = 0.26) y variaba de 26.1 a 50.2 cm dentro de rodales. La densidad promedio del pino ponderosa fue la mayor en la clase de 20.3–30.5 cm DAP y menor de las clases de 50.8–61.0 cm y >61.0 cm DAP. La densidad de árboles fue hasta 5.3 veces mayor que las consideradas típicas para bosques de pino ponderosa antes de la intervención para prevenir incendios. Las densidades de árboles muertos estuvo dentro del rango estimado para los bosques secos históricos del oriente de la cordillera Cascade, no obstante sólo 50% de los árboles muertos muestreados tuvieron un DAP >25.4 cm. Aunque se considera que los pájaros carpinteros cabeciblancos están estrechamente asociados con bosques primarios de pino ponderosa, mis resultados sugieren que podrían ser más adaptados a usar bosques donde predominan árboles de diámetro menor.     

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.        

Winter habitat selection patterns of Merriam's turkeys in the southern Black Hills, South Dakota

In northern areas of their expanded range, information on Merriam's turkeys ( Meleagris gallopavo merriami ) is lacking, specifically pertaining to wintering behavior and factors associated with winter habitat selection. Forest managers need detailed quantification of the effects of logging and other management practices on wintering habitats needed by Wild Turkeys and other wildlife. Therefore, we examined winter habitat selection patterns within ponderosa pine ( Pinus ponderosa ) forests and determined factors associated with use of farmsteads by Merriam's turkeys in the southern Black Hills, South Dakota. We radio-marked 86 female Merriam's turkeys (70 adults and 16 yearlings) and monitored them during winter (1 December–31 March), 2001–2004. Female Wild Turkeys used recently burned pine forest less than expected but selected farmsteads and stands of mature ponderosa pine ( 22.9 cm diameter at breast height [DBH] trees) for foraging sites. Within forests, female Wild Turkeys selected foraging sites with less understory vegetation and visual obstruction, and larger-diameter ponderosa pine. Ponderosa pine seed abundance varied among years, and pine seeds were most abundant in stands of 30–35 cm DBH with basal area of 22–28 m2 ? ha–1. Abundance of pine seeds may have influenced use of farmsteads by Wild Turkeys, more so than ambient temperatures or snow depth. In the southern Black Hills, management should emphasize open- to mid-canopy and mature-structural-stage pine stands, where seed production was greatest. During winters when mast from pine is unavailable, farmsteads likely provide nutritional supplementation and may be important for maintaining Merriam's turkey populations.     

Relative and seasonal abundance of three bark beetle predators (Coleoptera: Trogositidae, Cleridae) across an elevation gradient in ponderosa pine forests of north central Arizona

We examined abundance and flight periodicity of 3 predators of bark beetles (Coleoptera: Curculionidae, Scolytinae), Temnochila chlorodia (Mannerheim) (Coleoptera: Trogositidae), Enoclerus sphegeus (Fabricius) (Coleoptera: Cleridae), and E. lecontei (Wolcott) (Coleoptera: Cleridae), across an elevational gradient of ponderosa pine ( Pinus ponderosa Lawson) forests in north central Arizona. Predator populations were estimated at 10 sites in each of 3 elevation bands (low: 1600–1736 m; mid: 2058–2230 m; high: 2505–2651 m) for 3 years (2004–2006) using pheromone-baited funnel traps targeting 3 primary bark beetle species. We also investigated how predator abundance and flight seasonality related to those of 5 bark beetle species: Ips pini (Say), I. lecontei Swaine, Dendroctonus frontalis Zimmermann, D. brevicomis LeConte, and D. adjunctus Blandford. Temnochila chlorodia was most abundant in the low- and mid-elevation bands, whereas E. sphegeus was most abundant in the high-elevation band. Enoclerus lecontei showed no consistent elevational trend in abundance. Within each elevation band, changes in annual abundance of pooled predator species tracked shifts in abundance of pooled bark beetle species. In general, predator flight initiation coincided with or closely followed bark beetle flight initiation in the spring, but predator flight terminated before flight activity ended for most bark beetle species in the fall. In addition, the ratio of prey to predators was lowest in the summer and highest in the fall. This suggests that all bark beetle species examined may be provided temporal escape from their predators in the fall. For all 3 predator species, the pheromone-baited trap targeting D. brevicomis was less attractive than the pheromone-baited traps targeting I. pini and I. lecontei.     

Observations on woundfin spawning and growth in an outdoor experimental stream

The response of woundfin to different substrates and current speeds was investigated in an outdoor experimental stream. Fish spawned in groups of 15 – 20 over 5 – 10 cm rock substrate in a .06 to .09 m/sec current at a depth of 10 cm. Eggs were adhesive on the undersides of the rocks. Fish spawned under these conditions grew to approximately 55 – 60 mm TL in 5 months.     

Relationship between environmental and vegetational parameters for understory and open-area communities

Ten individuals from each of four tree species were selected, and their associated understory and adjacent open-area communities were sampled for both environmental and vegetational parameters, including light intensity, pH, litter depth, soil depth, and percentages of exposed rock, litter cover, living cover, shrubs, forbs, grasses, and annuals. The four tree species were ponderosa pine, Rocky Mountain juniper, Gambel oak, and snowbrush ceanothus. The study site was in the lower Uinta Mountains about 10 miles east of Kamas, Utah. Correlations among the various biotic and abiotic parameters were examined. The interplay of these factors in differentiating the understory and open-area communities is discussed.       

High rates of photosynthesis in the desert shrub Chrysothamnus nauseosus ssp. albicaulis

  Basic aspects of photosynthesis were investigated in white rubber rabbitbrush ( Chrysothamnus nauseosus (Pallas) Britt. ssp. albicaulis ), a common C 3 deciduous shrub native to arid regions of the western U.S. Under favorable field conditions, net photosynthesis (P n ) ranged from 36 to 73 mgCO 2 · dm – 2 · hr – 1 , which is relatively high for a woody species. The leaves from the actively growing flowering shoots exhibited higher P n than those on the vegetative shoots. P n also varied according to the age of the leaves and the location of the plants. P n did not light saturate even at quantum flux densities (QFD) equivalent to full sunlight. The light compensation point was relatively high (ca 100 μ mol · m – 2 · S – 1 ), perhaps due to the presence of a tomentose vestiture on the leaf surface. At high QFD's, the stomatal conductance was high (ca 520 mmol · m – 2 · s – 1 ) for a woody species. RUBP - carboxylase content of the leaves ranged from 20 to 22 mg per gram F.W., which is similar to that found in most C 3 crop species. These results suggest that rabbitbrush is able to maintain high rates of P n , at least under nonstressed conditions.           

Effects of long-term livestock grazing and habitat on understory vegetation

The herbaceous understory stratum contains most of the plant diversity in ponderosa pine ( Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.) forests of the American Southwest and provides critical food and habitat for many wildlife species. During the last century, this stratum has been affected by livestock grazing and by increased dominance of overstory trees. We sampled a unique grazing exclosure to examine the relative importance of long-term livestock grazing (grazed or ungrazed) and habitat (park or tree) on the understory community. We sampled 3 plots of 192 contiguous quadrats (each quadrat 0.5 m 2 ) in each of the 4 treatment combinations, for a total of 2304 quadrats. Species-area curves were generated by aggregating quadrats into nonoverlapping areas at grain sizes of 0.5 to 576 m 2 . The effects of habitat and grazing on species density were evident at very different scales. Species density was higher in park than tree plots at scales ≤32 m 2 but did not differ between habitats at larger scales. Species density differed minimally between grazed and ungrazed treatments at small grains, but grazed plots contained more species than ungrazed plots at larger grains. Grazing treatments differed at smaller grains (to 4–8 m 2 ) than did habitats (to 32 m 2 ), with respect to density of native species and graminoids. Grazed plots had more exotic species than ungrazed plots at all grain sizes, though few exotics were present. Twenty-two species were identified as indicator species associated with habitats and/or grazing treatments. Evaluations of plant community response to treatments would be improved by accounting for the grain at which data have been collected and analyzed and by identifying indicator species associated with various treatments. These data would enable more-informed conservation and management decisions.     

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 .     

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.     

Roost sites of the silver-haired bat ( Lasionycyeris noctivagans ) in the Black Hills, South Dakota

We investigated the roosting ecology of silver-haired bats ( Lasionycteris noctivagans ) in the Black Hills, of western South Dakota. Using radiotelemetry, we located 39 roosts, 10 of which were maternity aggregations containing 6 to 55 bats. The roosts were mostly in ponderosa pine ( Pinus ponderosa ) snags that averaged 39 cm diameter at breast height. Solitary bats preferred roosting under loose bark or in crevices in trees, regularly moving among trees. All maternity aggregations were found in tree cavities, primarily those created by woodpeckers. Roost trees were located in patches of forest with relatively high snag densities, about 21 snags/ha. This study suggests that snags play an important role in maintain silver-haired bat populations in ponderosa pine ecosystems.     

A comparison of the spherical densiometer and ocular methods of estimating canopy cover

Percent tree canopy cover in a ponderosa pine ( Pinus ponderosa ) forest of northeastern California was estimated by the point intercept spherical densiometer and ocular methods. Estimates derived by the two methods were similar (P > .05). The ocular method is recommended when understory vegetation is tall or clumped instead of randomly distributed, or if available field time limits sample size.     

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.     

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.        

Estimating root biomass and distribution after fire in a Great Basin woodland using cores and pits

Quantifying root biomass is critical to an estimation and understanding of ecosystem net primary production, biomass partitioning, and belowground competition. We compared 2 methods for determining root biomass: a new soil-coring technique and traditional excavation of quantitative pits. We conducted the study in an existing Joint Fire Sciences demonstration area in the central Great Basin. This area is representative of a shrub (sagebrush) ecosystem exhibiting tree (pinyon and juniper) encroachment. The demonstration area had a prescribed burn implemented 4 years prior to our study, and we sampled both control and burned plots. The samples were stratified across 3 microsites (interspace, under shrub, and under tree) and 4 soil depths (0–8, 8–23, 23–38, and 38–52 cm) to determine the effects of plant life form and burning on root biomass. We found that estimates of total root biomass were similar between quantitative pits and our soil cores. However, cores tended to show a more even distribution of root biomass across all microsites and depths than did pits. Overall, results indicated that root biomass differs significantly among microsites and soil depths and that the amount of root biomass at a given depth differs among microsites. Burning reduced root biomass in our study by 23% and altered the spatial distribution of root mass.     

Microhabitat affinities of Gambel oak seedlings

Previous work suggested that Gambel oak seedlings are rare in the northern parts of its range in Utah where summer rainfall is relatively low but should be abundant in southern parts of the range where summer rainfall is usually high. Gambel oak grades from a relatively minor component of a ponderosa pine/mixed conifer assemblage in the south to a virtually monotypic formation in the north, where it exists as long-lived clones.   Quadrat analysis in Arizona and New Mexico, within the oak zone, revealed a seedling density ranging from 120 to 1320 per hectare. We found a significant tendency of seedlings to be located on the NE (cool, shady) side of sheltering objects in the environment. Mature ponderosa pine ranged in density from ca 40 to 500 stems per hectare, whereas mature Gambel oak ranged from ca 10 to 20 genets per hectare with ca 1 to 7 ramets per clone. These results support our previous conclusion that Gambel oak in northern Utah probably became established as a minor component of a mixed pine/oak woodland at a time in mid-Holocene when summer rainfall was much higher than today.     

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.     

Standing crops and dynamics of phytomass and minerals in two salt desert shrub communities

Of two salt desert shrub communities studied in Curlew Valley, Utah, the Atriplex confertifolia- dominated community had 15% greater total midsummer phytomass than the Ceratoides lanata community. The larger Atriplex shrubs contained much more woody tissue for support of photosynthetic tissues than did Ceratoides. Atriplex aboveground phytomass and litter were about twice those of Ceratoides. Ceratoides litter was generally fine and easily decomposable, but Atriplex litter contained about equal proportions of coarse, resistant woody tissues and fine, easily decomposable material. Atriplex root phytomass was 1.3 times that of Ceratoides at the 2–30 cm depth, but at depths below 30 cm, Ceratoides exhibited up to three times greater root phytomass and had 23% more root mass overall. Net aboveground community primary production was estimated to be about one-third greater in the Atriplex than Ceratoides community. Turnover times for readily decomposable aboveground litter were quite similar, but, because Atriplex produced coarser litter, its overall rate was somewhat slower than that of Ceratoides. Analyses of selected minerals in plant parts, litter, and soil revealed that about 90% of the mineral capital is in the soil, mostly within organic matter. Nearly equivalent pools of mineral elements were found in the two communities, except for greater Na in the Atriplex community.     

Bottom-up factors influencing riparian willow recovery in Yellowstone National Park

After the elimination of wolves ( Canis lupis L.) in the 1920s, woody riparian plant communities on the northern range of Yellowstone National Park (YNP) declined an estimated 50%. After the reintroduction of wolves in 1995–1996, riparian willows ( Salix spp.) on YNP’s northern range showed significant growth for the first time since the 1920s. However, the pace of willow recovery has not been uniform. Some communities have exceeded 400 cm, while others are still at pre-1995 levels of <80 cm mean height. We took intensive, repeated measurements of abiotic factors, including soil and water-table characteristics, to determine whether these factors might be contributing to the varying pace of willow recovery. Willows at all of our study sites were “short” (<250 cm max. height) prior to 1995 and have recovered to varying degrees since. We contrasted “tall” (>250 cm max. height) willow sites where willows had escaped elk ( Cervus elaphus L.) browsing with “short” willow sites that could still be browsed. Unlike studies that manipulated willow height with fences and artificial dams, we examined sites that had natural growth differences in height since the reintroduction of wolves. Tall willow sites had greater water availability, more-rapid net soil nitrogen mineralization, greater snow depth, lower soil respiration rates, and cooler summer soil temperatures than nearby short willow sites. Most of these differences were measured both in herbaceous areas adjacent to the willow patches and in the willow patches themselves, suggesting that they were not effects of varying willow height recovery but were instead preexisting site differences that may have contributed to increased plant productivity. Our results agree with earlier studies in experimental plots which suggest that the varying pace of willow recovery has been influenced by abiotic limiting factors that interact with top-down reductions in willow browsing by elk.     

Effects of osmotic potential, potassium chloride, and sodium chloride on germination of greasewood ( Sarcobatus vermiculatus )

Greasewood ( Sarcobatus vermiculatus [Hook.] Torr.) (Chenopodiaceae) typically grows on salt-affected soils where its germination requirements may reflect characteristics necessary for establishment in saline environments. The objective of this study was to determine the effect of osmotic potential and specific ions on the germination of seeds from three populations of greasewood. Seeds were germinated at 20 C in solutions of polyethylene glycol with water potentials ranging from –0.3 to –2.2 MPa that contained 0 to 68480 µmol·L –1 sodium chloride (NaCl) or 0 to 53640 µmol·L –1 potassium chloride (KCl). Germination of two populations was reduced by increasing salt concentration and decreasing osmotic potential; germination of one population was reduced by declining osmotic potential. No seeds germinated at an osmotic potential lower than –1.6 MPa. For all populations, days to 50% of final germination increased and abnormal germination decreased as osmotic potential declined. Comparison of our results with those from other studies suggests geographic ecotypic development in response to osmotic potential and NaCl and KCl concentrations during germination.