number7 December 22, 1996

An information digest published by the USDA Forest Service, Rocky Mountain Research Station
Intermountain Fire Sciences Laboratory, P.O. Box 8089, Missoula, MT 59807

This issue of NUTCRACKER NOTES emphasizes current research and management in the genetics of whitebark pine. However, there are some articles that discuss cone crops, rust infections and prescribed burning. And, as a bonus, there are two interesting articles on limber pine ecosystems. You will also note some changes in the publication format and presentation of submitted articles. Your comments are graciously requested.

It snuck up on us, in a way. The sites on which whitebark pine grows are isolated, and the changes were not always dramatic. However, by the mid-1980s Steve Arno and others began to notice a significant decline in the stands. Eventually the signs pointed to three causal factors: the arrival of white pine blister rust, mountain pine beetle epidemics spreading up from lower-elevation lodgepole pine, and the suppression of fire which both reduced the number of high-elevation clearings that nutcrackers prefer as caching sites, and permitted subalpine fir to out compete whitebark pine.

At first, only scientists and academicians could afford to be concerned. Later, released from timber-based management by the newly-embraced ecosystem management, land managers in the national forests could explore ways of preserving existing stands and restoring those in decline. This problem is new: no one has managed or bred whitebark pine previously, so a genetically conservative strategy has been adopted. That strategy rests upon a time-tested theory about adaptation: in the absence of genetic tests, use local seed for reforestation. The genetic tests in question are tests of adaptive variation, presently being carried out by the Intermountain Station. Once the information is available we will know how far we can safely move seed without the seedlings suffering maladaptation.

If the decline were simply the result of mountain pine beetle attacks or fire suppression, the strategy would be simpler. The important thing to remember, however, is that white pine blister rust is now a permanent feature of North American forests. At one time, presumably, it was as devastating in the Eurasian forests where it evolved. However, the rust and the Eurasian hosts have also presumably evolved together to permit both to survive and reproduce. Eventually, this should occur in North America as well, and in fact a "mass selection" program is underway as the rust identifies susceptible individuals and eliminates them.

Geneticists at the Intermountain Lab seek to both imitate and speed evolution in two ways: by facilitating the process of mass selection and by developing a breeding program similar to that used in western white pine. Ray Hoff’s silvicultural guidelines for whitebark stands will promote a large supply of naturally regenerated seedlings for the rust to test. Hoff has already been pleasantly surprised by the percentage of resistant seedlings coming from stands that have experienced over 90% mortality by the rust. As time passes, this percentage should increase. A breeding program such as the one established by Bingham for western white pines consists of identifying trees that carry resistance genes and breeding them, thus making combinations possible that would not have occurred otherwise. Unfortunately, considering the delayed sexual maturity of whitebark pine, generation times are extremely slow. A successful breeding program may condense the work of evolution from thousands of years to a century or two (forest genetics is not a field for those looking for quick results! Might I suggest Drosophila melanogaster...)

At the same time, we recognize the larger problem of conservation of the whole species. The death and destruction we see in the stands prompts a natural response: we must save whatever is left, on all the affected acres! We have absorbed the concept, established by Aldo Leopold’s Sand County Almanac, that we must save all the pieces. We look at the example of modern agricultural breeders as they painfully try to reconstruct the ancestral variability of such species as wheat and corn. This is unmistakably a vulnerable position which we must seek to avoid. But how much of the species must we conserve?

Our impulse to save everything runs headlong into the realities of budget, time, and people. This is why genetic studies are critical. In order for managers to be able to make realistic decisions about what to conserve and why, we need an understanding of whitebark’s genetic variability, and how it is distributed (this is termed "genetic structure").

Every species has a certain amount of genetic variability. Genetic variability is important because it influences both an individual’s short-term fitness and a species’ long-term adaptability to changing environmental conditions. A species’ total genetic variability is divided either within individuals, between individuals within a population, or between populations.

High genetic variability within populations means that there are many different "versions" of genes among the different trees in the population. Low variability, on the other hand, would indicate only a few different "versions." (These "versions," called alleles, are simply alternate forms of a gene).

High genetic variability between populations, on the other hand, would indicate that populations that are separated from each other are genetically different. One common way that conifer populations differ is that those from northern latitudes and higher elevations are often more cold tolerant than those from southern latitudes and lower elevations. Low genetic variability would mean that populations separated from each other are not that different from each other. For instance, a western white pine from the Clearwater National Forest may be genetically more different from another tree standing 100 yards away than it is from a tree in western Washington.

Typically, pines have high variability within populations. In some cases, there is high variability between populations; this usually indicates that a species is tightly adapted to specific site conditions. Douglas-fir is a good example of one such species, which is why seed transfer guidelines for DF are rather specific. Low variability between populations, on the other hand, indicate a species that is more broadly adapted and will do well on a variety of sites. Western white pine is such a species.

There are significant implications to finding species with low variability between populations. What it means, in effect, is that one population is not much different from another, so fewer populations would need to be preserved to adequately capture genetic variability. An additional bonus is that seed transfer guidelines may be relatively loose. On the other hand, finding species with high variability between populations means that more collections must be made and seed transfer guidelines must be relatively tight.

It is difficult to predict the genetic structure of a species. In a forthcoming paper in the American Journal of Botany, Jerry Rehfeldt supports the theory that "genetic structure, first and foremost, reflects the uncertain, chance events that are interspersed throughout evolutionary history." The unusual life history of whitebark pine, especially the animal-mediated dispersal of its seeds, may suggest an unusual genetic structure. Studies are underway to find this out.

Once we know how different one population is from another, how do we conserve? Do we establish in situ reserves? Essentially, many whitebark pines are already in such reserves by virtue of their location in wilderness areas, national parks, and other areas with difficult access. This is patently not the answer if the stands are simultaneously threatened by blister rust, mountain pine beetle, and encroachment. It may be an answer, however, if we can actively manage for whitebark pine.

We can also establish ex situ reserves by collecting seeds and either holding them in storage or growing them and establishing them in clonebanks. Long-term storage is probably not an acceptable solution as seed viability will be lost over time and collections will be difficult to replace. A system of clonebanks, however, may be a good solution.

A combined approach may be to first, establish the genetic structure of the species and use that to decide how many populations must be conserved. Second, choose and manage those populations as in situ reserves by following Hoff’s silvicultural recommendations (thin out the subalpine fir and encourage continuous regeneration of whitebark pine). This will also encourage the rust to test the seedlings and eliminate those that are not resistant. To whatever extent is possible, the stands should also be protected from mountain pine beetle by managing the lower-elevation lodgepole. Since bark beetles prefer large, old trees, a regular supply of regeneration should also allow for young trees to grow in and replace them. Third, seed collections should be made from those populations. Fourth, the collected seeds should be used to establish several clonebanks and manage them the same way as the in situ reserves, by keeping the subalpine fir down and encouraging constant regeneration.

Dr. Donna Dekker
USDA Forest Service
Intermountain Research Station
Forestry Sciences Laboratory
Moscow, ID 83843
Phone: 208-883-2324
Email: d.dekker:s22l04a

Whitebark Pine in the Lower Salmon River Country -- Linking field monitoring with cyberspace by Marcy Mahr, Research Biologist, Craighead Wildlife-Wildlands Institute, 5200 Upper Miller Creek Rd., Missoula, MT 59803 (406) 251-3867

This summer six field biologists from the Craighead Wildlife-Wildlands Institute (CWWI) collected detailed ecological data on whitebark pine stands as part of our broader habitat study of the Salmon-Selway Ecosystem. Whitebark pine, a key grizzly bear food, was dominant or co-dominant (with at least 10% relative canopy cover) in 5% of our sampled plots. Tenth-acre plots were located in polygons delineated on 1:24000 scale quadrangles, spectral maps produced from satellite imagery. Sampling, or ground-truthing, extended west from the Bighorn Crags of the Salmon National Forest to the South Fork of the Salmon River and the McCall area, and south to the Sawtooth NRA.

CWWI's field methods were initially based on the whitebark monitoring protocol developed by Kate Kendall in Glacier National Park. Meeting the criteria of a minimum of 20 whitebark individuals was easily achieved at all sites; some stands were so dense with a subcanopy of whitebark we modified our strategy to sample a maximum of 15 individuals per size class to increase our efficiency. All mature trees (>5 in dbh) in the plots were sampled because we did not encounter >15 individuals in pole, medium or large tree classes. Our data forms resemble USFS Region 1 ECODATA general, location, plant composition, and tree data forms.

Our whitebark pine data awaits analysis yet preliminary inspection show very low blister rust infection and mortality rates. Moreover, there was a noticeable absence of Ribes spp. in or proximate to our sample sites. We are intrigued by the contrast of whitebark pine habitats in the rugged terrain south of the main Salmon River with the mesic, glacially- sculpted habitats of Glacier National Park where avalanche chutes teem with lush growth and encourage Ribes spp. to grow adjacent to whitebark stands. We also observed good cone production at several sites and hope to incorporate more rigorous methods of assessing production into next summer's research.

CWWI, a multidisciplinary research center, integrates field-based ecological research, innovative uses of technology, and scientific activism. For more information please contact: Marcy Mahr, Research Biologist, Craighead Wildlife-Wildlands Institute, 5200 Upper Miller Creek Rd., Missoula, MT 59803 (406) 251-3867.

The Rocky Mountain Front: A Landscape in Transition by Dayna Ayers Baumeister, Training Web Fellow, Division of Biological Sciences, University of Montana, Missoula, MT

In northcentral Montana, running north to south along the eastern fringe of the Rocky Mountains there lies a landscape unlike any other in the West. Here the sweeping Great Plains and the majestic Rocky Mountains collide, creating a diverse transition zone that appears frozen in time. Low human occupation has left most of this landscape undeveloped and human impacts have been slight. Recent evidence, however, has shown that this landscape has actually changed dramatically since first settlement by Euro-Americans in the 1860s. Over the last 130 years, the landscape of the Front has shifted from what once was a homogenous fescue prairie to a more diverse landscape, dominated by patches of limber pine (Pinus flexilis) and small aspen (Populus tremuloides) and shrub communities.

Environmental factors leading to these changes are diverse and often complex. Historically, an array of ecological processes, including frequent fires and bison grazing successfully prevented the invasion of tree and shrub communities into the grasslands. For several millennia, the disturbance regime of grazing and fire restricted non-grassland communities—particularly limber pine—to bare, rocky outcrops, cliffs, and steep riparian corridors—areas absent of these disturbances. In fact, these areas are the only places on the Front where one can find limber pine older than 150 years.

The most recent invasion of limber pine into the grasslands on the Rocky Mountain Front probably began around Euro-American settlement in the 1860s. With the suppression of fires and the elimination of bison grazing, conditions on the Front became more favorable for limber pine and other tree and shrub communities to establish. Limber pine seedlings were now able to survive long enough to reach maturity and produce cone crops. Research has shown that most large limber pine on the Front today are between 90 and 120 years old, established during this time of intensive grazing and land settlement.

These changes suggest that gradually, over the last century, fewer fires and year-long grazing allowed for successful expansion of limber pine—from river corridors and rocky outcrops into the grasslands. As limber pine spread eastward, the conditions established by their presence created new habitat for a diversity of wildlife species and probably accelerated the rate of limber pine invasion. Forested areas are known to ameliorate harsh winds and trap drifting snow, yielding greater soil moistures throughout spring and summer. This probably increased the rate of successful germination of limber pine and promoted a diversity of understory plants. On the fringes of these limber pine stands, aspen groves and shrub communities have established, subsequently adding to the diversity of the landscape. This diversity, in turn, has provided additional habitats for wildlife on the Front.

Wildlife biologists and land managers are only now beginning to understand the relationships of limber pine and associated wildlife communities on the Front. The importance of limber pine as habitat differs among species; the relationships can be grouped into three general categories: incidental, suitable, and exclusive/obligatory. Incidental relationships include those with species such as grizzly bear, mountain lion, and elk—species which do not necessarily depend on limber pine but use this habitat frequently for feeding, cover or shelter. Wildlife species that find limber pine suitable habitat include mule deer, mountain chickadee, and chipping sparrow. These species can be found in other habitats on the Front, but prefer areas with limber pine.

A few species on the Front are almost exclusively found in limber pine, including blue grouse, Townsend’s Solitaire and Clark’s nutcracker. The Clark’s nutcracker, in fact, has evolved into a obligatory relationship with limber pine. The nutcracker feeds exclusively on the large, nutritious seeds. These corvids open cones while they are still on the tree, store up to 125 seeds in their pouch, and then cache them in groups of two to a dozen in sites that are accessible during winter. Clark’s nutcracker caches between 70-90% of the annual seed production of limber pine, and often caches are unrecovered; the seeds subsequently can develop into seedlings. A similar symbiotic relationship occurs between the Clark’s nutcracker and other five-needled, stone pines, including white bark pine (Pinus albicaulis).

As local residents and visitors to the Front today can attest, the landscape is diversified by its many habitats. The ridgetops and hillsides support expanding stands of limber pine. These stands are lined with patches of shrubs such as rose, serviceberry, shrubby cinquefoil, and chokecherry. In the moister ravines, aspen groves grow thick and provide a cool microclimate during hot summer months. The patchwork of these different types of vegetative communities supports an array of wildlife species including elk, mule deer, grizzly bear, wolves, coyotes, ruffed grouse, white tailed deer, Bohemian waxwing, mountain bluebird, Northern flicker, mountain lions and badgers, among others. The increased habitat diversity of the Front today supports both higher populations and a greater diversity of wildlife species than recorded by settlers 130 years ago.

The environmental conditions of the Front, as we understand them today, suggest that limber pine has become an integral component of the landscape, both by defining new habitats and supporting wildlife. But this will not continue into perpetuity; the landscape is still changing. Ecological processes are shifting yet again, and limber pine appears to be dying out. Since the early 1970s, an infestation of blister rust has infected many stands along the Front, gradually killing many trees and often leaving behind "ghost" stands. These foreboding changes may be accelerated by land use practices, climate change or possibly new, yet unknown, inhospitable conditions. Fortunately, the extent of the losses have not yet reached those experienced by limber pine’s cousin, the white bark pine. Research is currently underway examining the magnitude of blister rust infection in limber pine. (Please see the related article, "Status of Limber Pine stands of the lower and upper timberline zones in the Northern Rocky Mountains," pp. xx-xx, this issue.)

Without fully understanding the ecological processes influencing the current distribution of limber pine and the subsequent affects of these stands on the landscape ecology of the Front, it will be difficult to ascertain what the future environment of the Front will be. Quite possibly in another 130 years, biologists will describe the landscape of the Rocky Mountain Front, and hypothesize on the ecological processes that caused the shift from a savanna landscape of limber pine foothills with a diverse array of wildlife to the present landscape, one perhaps of homogenous, thick conifer forests, or perhaps maybe once again, of vast prairies of fescue grassland, treeless and bare, except for a scattered few, old limber pine and the occasional bull elk.

Shoshone National Forest Whitebark Pine Cone Surveys by Mark Hinschberge, Shoshone National Forest, 808 Meadow Lane, Cody, WY 82414

The Shoshone National Forest has participated in the whitebark pine cone surveys since 1980. Now the forest has 4 transects that are surveyed for cone production for the Interagency Grizzly Bear Study Team. These transects are C (Republic Creek), D (Upper Sunlight), H (Moccasin Basin), and U (Brent Creek). This year's cone production was higher than the past several years. Transects C and D, which in 1995 had no cones on any of the 20 trees, produced 157 (15.7 mean cones per tree) and 235 cones (23.5 mean cones per tree), respectively. Transects H and U produced 22.4 and 29.8 cones per tree, respectively. The Brent Creek transect has exceeded 22 cones per tree each year since 1992, however the Moccasin Basin site has not exceed 20 cones per tree since 1991.

The production on the Shoshone National Forest should be considered good when compared with the entire ecosystem. Throughout the ecosystem the mean cones per tree averaged 24.7. Not since 1989 has the average cones per tree exceeded 20. According to Mattson, Blanchard and Knight (1992. J. Wildl. Manage. 56(3):432-442) widespread use of whitebark pine seeds by grizzly bears generally occurs when production exceeds 22 cones per tree. They have also found that when cone production is high, the number of nuisance bear management actions and grizzly bear mortalities is low. So there should have been less bear relocations and mortalities this past fall on the Shoshone and in the entire ecosystem than in the fall of 1995.

Jim Hadfield, Forest Pathologist, and Paul Flanagan, Forest Entomologist
Wenatchee National Forest
Wenatchee, Washington

Ann Camp, Research Silviculturist
Wenatchee Forestry Sciences Laboratory
Wenatchee, Washington

In 1996 we began a whitebark pine mortality survey in the eastern Washington Cascades. The objective of the survey is to determine whitebark pine mortality levels and identify mortality causal agents. The survey is done by examining all whitebark pines within 5 to 10 feet of both sides of a compass line extending through locations containing whitebark pine. Transect width depends upon apparent stocking in the strands. A minimum of 50 whitebark pines are examined on each transect. Trees are classifies into size classes of seedlings (<1" DBH), saplings (1.0"-4.9"), poles (5.0"-8.9"), and mature (9+"). Trees are classified as alive, recent dead (<5 years), old dead (5-10 years), and very old dead (>10 years). The presence of white pine blister rust infections is recorded and infections are classed as lethal or nonlethal. Presence of insects is recorded if the surveyors believe insects are responsible for tree death or are active in living and dying trees. Other damaging affects seen on the trees are recorded, typically these include porcupine feeding, other rodent feeding, bear feeding, mechanical damage, fire damage, and brown felt foliage disease.

Six sites were surveyed in 1996. Mortality data from these sites are shown in Table 1.

Overall, 8.4 percent of the trees were dead, and only 1.5 percent of the trees died within the last five years. Assigning causes of mortality is difficult in many cases, especially on trees classed as very old dead.

White pine blister rust infections have been found at all sites surveyed (Table 2). The infections have not been aged, but historical reports indicate white pine blister has been present in the eastern Washington Cascades for more than 60 years. The percent trees shown infected by white pine blister rust is conservative. Most of the pole and mature size trees occur in clumps and are quite limby making close examination difficult. Many trees have had bark gnawed by porcupines and other rodents, most likely squirrels. Some, but by no means all, of the gnawing is associated with blister rust infections. We are not able to make accurate identification of blister rust infections on old dead and very old dead trees.

Many trees have been killed by fires, especially on the Wenatchee National Forest.

We plan to survey several more eastern Washington Cascade Mountain whitebark pine stands for mortality in 1997.

Genetic Relationships Associated with Krummholz and Stem-cluster Growth Forms of Pinus albicaulis in the Eastern Sierra Nevada, California by Constance I. Millar and Deborah L. Rogers, Pacific Southwest Research Station, USDA Forest Service, P.O. Box 245, Berkeley, CA 94701

Whitebark pine (Pinus albicaulis Engelm.) possesses several attributes, atypical of pines in general, that putatively affect its local, spatial, genetic structure. It is one of only 20 species of pine worldwide (approximately 100 species within the genus Pinus) with wingless, animal-dispersed seeds (Tomback et al. 1990). This method of seed dispersal has significant implications for the regeneration capacity of the species, its ability to colonize in new locations (Tomback 1982), and the genetic structure of the species (Furnier et al. 1987). It often grows in multi-stemmed clusters, an inferred consequence of seed-cache origin (Tomback and Linhart 1990). Furthermore, the species has the potential to reproduce vegetatively, with unmeasured impacts on genetic structure. These features are responsible for the different growth forms in the species — a high elevation krummholz thicket form (possibly clonal), and a multiple-stem form (occasionally single stems) observed mostly at lower elevations or sheltered habitats. Both growth forms are observed in the eastern Sierra Nevada of California.

The mutualistic relationship between the Clark's nutcracker (Nucifraga columbiana) and whitebark pine has been well studied (e.g., Tomback 1982, Tomback et al. 1990). Several studies have investigated the genetic implications for this method of seed dispersal in the multi-stem cluster form in the Rocky Mountains in Alberta (Furnier et al. 1987) and in Wyoming (Linhart and Tomback 1985). However, the effects of vegetative reproduction, genetic relationships with elevation, and range-wide genetic variation have not been well studied. Interaction among these variables could have considerable impact on the genetic structure of whitebark pine. For example, both caching activity and tendency to reproduce vegetatively could be affected by elevation.

The eastern Sierra Nevada presents an ideal venue in which to explore some of these genecological relationships. The western and especially southwestern extension of the species' range in the Sierra Nevada/Cascades have received little scholarly attention. Typically, coniferous species with a similar span in their natural range exhibit genetic differentiation between Sierra Nevada and Rocky Mountain populations, suggesting different genetic architecture in whitebark pine and potentially different ecological behavior of nutcrackers. A range-wide monoterpene study of whitebark pine provides evidence of genetic differentiation between Californian populations and the rest of the species' range (Zavarin et al. 1991). We undertook a genetic study of whitebark in the eastern Sierra Nevada specifically:

Three regions, each with two contrasting occurrences of whitebark pine — stem clusters at lower elevation and krummholz patches at higher elevation — were selected in the eastern Sierra. Foliar samples were collected at each of the six sites and studied with allozyme analysis. Every stem in each selected stem-cluster was sampled, while evenly-spaced samples were collected from the perimeter of every selected krummholz thicket. Results from one of the three regions have been statistically analyzed. These results suggest that stem-clusters at lower elevations have a lower proportion of clones (i.e.,distinct genotypes) per group and closer genetic relationships among the clones present in each cluster than do the higher-elevation krummholz thickets. However, using measures of expected heterozygosity, polymorphic loci, and numbers of alleles per locus there is little difference between the krummholz and stem-cluster forms in this one region. Thus, the genetic differences are in distribution of variation rather than amount.

One contributing factor to the differences in genetic structure observed could be that the stem-clusters may be established by only one or a few caching events, probably in the same season, while the krummholz thickets could be the result of multiple, multi-year caching events. Krummholz patches might grow in dimensions over time by gradually recruiting seedlings from new caches, these seedlings having a better chance for survival than those not sheltered by an existing cache. (Isolated single stems or seedlings in the upper elevation areas were not observed.)

Overall, the genetic diversity, as measured by expected heterozygosity and percent polymorphic loci, is in the mid-range for coniferous species in California, and similar to values reported for Chamaecyparis lawsoniana, Cupressus macrocarpa, and Calocedrus decurrens. Values for observed and expected heterozygosity are at the low end of the range reported for disjunct populations of whitebark pine in and near the Great Basin (Yandell 1993). This is interesting as one might have expected the disjunct populations to show lower heterozygosity due to inbreeding, a consequence of their geographic isolation. The suite of enzymes used in the 1993 study, however, is not identical to the one used in this study.

We also intensively sampled one of the krummholz patches in one of the three study regions to help:

Analysis of the allozyme patterns in the remaining two regions is ongoing. Once completed, this study will contribute to the growing awareness of the relationship between genetic variation and ecological conditions in whitebark pine, together with other current genetic studies of the species (e.g., those by D. Dekker-Robertson and D. Tomback).

Furnier, G.R., P.Knowles, M.A. Clyde, and B.P. Dancik. 1987. Effects of avian seed dispersal on the genetic structure of whitebark pine populations. Evolution 41: 607-612.

Linhart, Y.B. and D.F. Tomback. 1985. Seed dispersal by nutcrackers causes multi-trunk growth form in pines. Oecologia 67: 107-110.

Tomback, D.F. 1982. Dispersal of whitebark pine seeds by Clark's nutcracker: A mutualism hypothesis. J. Animal Ecology 51: 451-467.

Tomback, D.F., L.A. Hoffman, and S.K. Sund. 1990. Coevolution of whitebark pine and nutcrackers: implications for forest regeneration. In Proceedings - Whitebark pine Ecosystems: Ecology and Management of a High Mountain Resource, U.S.D.A. Forest Service, Intermountain Research and Experiment Station, Ogden, Utah, pp 118-129.

Tomback, D.F. and Y.B. Linhart. 1990. The evolution of bird-dispersed pines. Evol. Ecology 4: 185-219.

Yandell, U.G. 1993. An allozyme analysis of whitebark pine (Pinus albicaulis Engl.) M.S. thesis, University of Nevada, 63 pp.

Zavarin, E., Z. Rafii, LG. Cool, and K. Snajberk. 1991. Geographic monoterpene variability of Pinus albicaulis. Bio. Syst. Ecol. 19(2): 147-156.

Population Genetics of Whitebark Pine in the Canadian Rockies by G. Jon Stuart-Smith and S. Ellen Macdonald, Department of Renewable Resources, 2-30D Earth Sciences Building, University of Alberta, Edmonton, AB T6G 2E3, Phone: 403-492-4155, gjs@gpu.srv.ualberta.ca

Research on whitebark pine has been focused on populations in Idaho and Montana. However, besides some insightful worked done on the genetic structure of tree clumps (Furnier et al 1987), little is know about how whitebark pine ecology differs in the Canadian Rockies compared to populations further south. As part of the requirements of a Masters of Science degree at the University of Alberta (GJS-S), we propose to examine the population dynamics and population genetics of whitebark pine in the Canadian Rockies.

Whitebark pine does not form climax communities in the Canadian Rockies such as are found further south (Arno and Hoff 1989). For these populations, the importance of fire and other disturbance types for regeneration and what effect fire suppression has had on this regeneration has not been examined. By examining age distribution within stands in relation to evidence of disturbance, we should be able to determine the dependence of whitebark pine on fire for regeneration. Stands of individuals approximately the same age would indicate that whitebark pine requires fire to remove competition. In contrast, if stands are composed of individuals of all ages, then regeneration is occurring continuously, and factors other than fire are more important in creating the disturbance necessary for regeneration.

Recent studies in Northwestern Montana attributed a decline in whitebark pine populations to successional replacement (by the more shade tolerant conifers Engelmann spruce and subalpine fir) resulting from fire suppression (Keane and Arno, 1993; Keane et al. 1994). Successional replacement will increase the average age of individuals by reducing regeneration. White pine blister rust and mountain pine beetle have also been implicated in the decline of whitebark pine in Western Montana (Keane and Arno 1993, Tomback et al 1995).

By examining population origin and age structure in relation to disturbance, successional replacement, and blister rust and beetle infections, we will be able to infer the importance of disturbance in establishment and the role of fire suppression, insects and disease in causing any decline in whitebark pine.

Trees, conifers specifically, have high genetic variation within species and within populations but relatively low genetic differentiation among populations (Loveless and Hamrick 1984). Several life history traits associated with this pattern of variability are: long-lived, large geographic range, outcrossing and wind pollination. Nutcracker seed dispersal results in populations with highly related clumps of individuals but little or no relation between clumps (Furnier et al. 1987). Therefore, although whitebark pine is outcrossing and wind pollinated, animal dispersal may decrease gene flow among populations resulting in greater population differentiation.

There have been no comprehensive studies of genetic variation in whitebark pine (Brussard 1990). Studies on three widespread species of Eurasian stone pine showed high genetic variation at the species level but low variation among populations (Politov et al. 1994). In contrast, Schuster et al. (1989) found high genetic differentiation between two, more isolated, populations of limber pine in Colorado.

Genetic processes such as genetic drift and founder effects are likely to result from the unique reproductive ecology of whitebark pine. Founder effects and genetic drift both result in increased differentiation among populations (Carson and Templeton 1984). Inbreeding, caused by isolated populations and low gene flow, may further increase differentiation among populations. Furnier and Dancik (1992) suggest that inbreeding is high in whitebark pine. We hypothesize that the genetic effects of isolation, restricted gene flow, and small population size on whitebark pine found in the Canadian Rockies may have resulted in low within-species and within-population genetic variation, compared to more widespread conifer species, and a higher portion of the genetic variation being partitioned among populations.

We propose to examine stand structure and disturbance type for sites from Waterton National Park in southern Alberta, through the four Rocky Mountain National Parks of Banff, Jasper, Kootenay and Yoho, to the limit of the range of whitebark pine in the Willmore Wilderness area in Central Alberta. These sites will then be revisited in the fall to collect seed for allozyme analysis. We hypothesize that fire disturbance has played an important role in the regeneration of whitebark pine and that fire suppression and successional replacement have reduced the size and increased the age of whitebark pine populations in the Canadian Rockies. We also expect that isolation, reduced gene flow and bird mediated seed dispersal have resulted in a low species genetic variation with the majority of that variation being among populations. Therefore, examination of population age and size and genetic variation will indicate how well equipped whitebark pine is to deal with future change such as global warming or disease epidemics and verify the need for an active management approach to ensure the continued contribution of whitebark pine to the subalpine ecosystem.

Arno, S.F. and R. J. Hoff. 1989. Silvics of Whitebark Pine (Pinus albicaulis). USDA For. Ser. Gen. Tech. Rep. INT-253. 11 pp.

Brussard, P.F. 1990. The role of genetic diversity in whitebark pine conservation. In: Schmidt, W.C. and K. J. McDonald (comps). Proceedings - Symposium on whitebark pine ecosystems: ecology and management of a high-mountain resource; 1989 Mar 29-31; Bozeman, MT. USDA For. Serv. Gen. Tech. Rep. INT-270.

Carson, H.L. and A.R. Templeton. 1984. Genetic revolutions in relation to speciation phenomena: The founding of new populations. Ann. Rev. Ecol. Syst. 15:97-131.

Furnier, G.R. and B.P. Dancik. 1992. Inbreeding in natural populations of whitebark pine. In: Proceedings, 12th North American Forest Biology Workshop, The role of physiology and genetics in forest ecosystem research and monitoring, Sault Ste. Marie, Ont., Canada. Ontario Ministry of Natural Resources. p161.

Furnier, G.R., P. Knowles, M.A. Clyde and B.P. Dancik. 1987. Effects of avian seed dispersal on the genetic structure of whitebark pine populations. Evolution. 41(3):607-612.

Keane, R.E. and S.F. Arno. 1993. Rapid decline of Whitebark Pine in Western Montana: evidence from 20-year remeasurements. West. J. Appl. For. 8(2):44-47.

Tomback, D.F., J.K. Clary, J. Koehler, R.J. Hoff and S.F. Arno. 1995. The effects of blister rust on post-fire regeneration of whitebark pine: The Sundance burn of Northern Idaho (U.S.A.). Conserv. Biology. 9(3):654-664.

Keane, R.E., P. Morgan, and J.P. Menakis. 1994. Landscape assessment of the decline of Whitebark Pine (Pinus albicaulis) in the Bob Marshall Wilderness Complex, Montana, USA. NW Sci. 68(3):213-229.

Loveless, M.D. and J.L. Hamrick. 1984. Ecological determinants of genetic structure in plant populations. Ann. Rev. Ecol. Syst. 15:65-95.

Politov, D. V. and K. V. Krutovskii. 1994. Allozyme polymorphism, heterozygosity, and mating system of stone pines. In: Schmidt, W.C. and F.K. Holtmeier (comps.). Proceedings - international workshop on subalpine stone pines and their environment: the status of our knowledge; 1992 Sept 5-11; St. Moritz, Switzerland. USDA For. Ser. Gen. Tech. Rep. INT-GTR-309.

Schuster, W.S., D.L. Alles, and J.B. Mitton. 1989. Gene flow in Limber pine: evidence from pollination phenology and genetic differentiation along and elevational transect. Amer. J. Bot.76(9):1395-1403.

Population Dynamics of Whitebark Pine Forests in the South Warner Wilderness, Northeastern California by Pete Figura, Department of Biological Sciences, Humboldt State University, Arcata, CA 95521, Phone: 707-826-3346, pjfl@axe.humboldt.edu

Although the development of management strategies for ensuring the survival of whitebark pine and its valuable seed crops depends on a thorough knowledge of the species' habitat, competitive relationships, and patterns of stand development and regeneration, relatively little is known about its stand dynamics, especially within lower-elevation stands where it competes with other tree species (Mattson and Reinhart 1990). This lack of knowledge appears to be particularly acute for stands in the southwestern portion of the species' range, as most ecological research on the species has been conducted in the northern Rocky Mountains. Because of this gap in our knowledge and based on the preliminary field work and observations of Gregg Riegel (USFS PNW Research Station), Dale Thornburgh, and John Sawyer (Humboldt State University), I began an investigation of the stand dynamics of whitebark pine-dominated forests in the South Warner Wilderness in summer, 1994. The primary objective of this project is to inventory the age structures and other structural characteristics of these forests across their elevational range in order to 1) determine the overall health and the current regenerative status of whitebark pine in these mountains, and 2) to better understand the primary physiographic and disturbance-related factors that affect forest development in these stands.

The Warner Mountains are part of the western edge of the Great Basin Province-the western slopes of the range drain into the Pit River (tributary to the Sacramento River) and the eastern slopes drain into landlocked basins. Structurally the range is characteristic of the Great Basin, although its rock is compositionally related to the Modoc Plateau (Macdonald and Gay 1966). The rocks comprising the range are typically Miocene volcanic rocks overlying Oligocene sedimentary rocks (Duffield and Weldin 1976).

The study area comprises the upper portion of the western slope of the South Warner Mountains in the vicinity of their highest peak and contains one of the larger areas of well-developed, relatively contiguous whitebark-dominated stands in the range. While most environmental variable (slope, aspect, etc.) are relatively consistent within the area, elevation varies considerably. The eastern boundary of the study area follows the crest of the range northward for approximately 1.3 miles. The western boundary follows the 2280 m contour along the western slope. The distance from the 2280 m contour to the crest of the range (the east-west study area length) averages approximately 1.6 miles.

Whitebark pine occurs as the dominant or co-dominant species in extensive stands in the Warner Mountains between approximately 2280 m and the crest of the range (to 3043 m). These stands typically consist of pure whitebark pine above approximately 2560 m and whitebark pine and white fir below 2560 m. Lodgepole pine and western white pine are very infrequent associates. Below 2280 m white fir dominates the forests, with Jeffrey pine, Washoe pine, and western juniper as common associates. Scattered whitebark pines occur down to approximately 2000 m. Riegel et al. (1990) identified three whitebark-dominated habitat types in South Warner range and found that the composition and structural features of these types vary in response to changes in elevation and aspect.

Field work. I established permanent plots (8 x 50 m) at 91 m (300 ft) elevation intervals along six systematically-located transects, which ran from 2280 m to the crest of the mountains (between 2835-3043 m within the study area). The transects were located approximately 412 m apart. Seven or eight plots (depending on ridgetop elevation) were established along each transect for a total of 44 plots. Environmental variables inventoried at each plot included elevation, aspect, slope angle, slope configuration, topographic position, litter depth, amount of surface rock and down wood, habitat type, livestock grazing sign, and fire evidence. Vegetation was inventoried by assigning cover class values to the dominant vascular plant species in each growth form. Variables recorded for coniferous stems >1.4 m tall included stem density, height, diameter at breast height, and presence/absence of white pine blister rust and dwarf mistletoe. All single-stemmed individuals and the largest sound stem of multi-stemmed individuals (901 stems total) were cored at approximately 0.3 m above ground level with an increment borer. Stems were tagged with numbered aluminum tags (1999 total stems). Density, basal diameter, height, and health were also recorded for stems between 0.5 and 1.4 m tall. Numbers of seedlings (<0.5 m) were tallied within an 8 x 25 m subplot. Whether each stem and seedling occurred individually or was part of a multi-stemmed cluster was also recorded. Additionally, I cored each stem of ten multi-stemmed clusters (elevation approximately 2500 m) to determine whether each stem in multi-stemmed clusters was the same age or nearly the same age as other stems in the cluster (sampled clusters averaged 3 stems; 29 cores total).

Lab Work and Data Analysis. Cores were mounted, sanded, and tree ages at coring height were determined with a dissecting microscope. Current work involves analysis of the age and structural data. I plan to primarily compare the effects of elevational differences on forest structure, age distribution, and stand development patterns. Data may also be analyzed to determine if age and structural differences occur based on slope aspect (although the aspects were fairly consistent from plot to plot) and if each of the different whitebark-dominated habitat types show unique structural and age characteristics. Age distributions will also be evaluated in terms of historic grazing intensity and what is known of historic fire occurrence.

The whitebark-dominated stands within the study area appear relatively healthy. Regeneration is occurring across all elevations and habitat types. Although currently present, the incidence of white pine blister rust is low (probably less than 2-3% trees infected) and seems to be primarily relegated to areas below approximately 2650 m. Annual mortality was less than 1% in 1991 (327 trees) and 1993 (539 trees) (Riegel and others, unpublished data), although Ribes cereum and/or R. montigenum occur throughout much of the study site. Perhaps the region's somewhat arid climate and the relatively open stand structure of the Warner range are factors helping to maintain white pine blister rust in an incipient rather than epidemic sate. Whitebark pine (and white fir) appear to be expanding into Artemisia-Wyethia and some Populus tremuloides-dominated areas, based on the preliminary field observation of smaller size-class trees scattered about the "sage steppe" areas near the forest-sage interfaces. This invasion may be a result of the complex interaction of grazing practices, fire suppression, and climate. Vale (1975, 1977) suggested that historic grazing practices were responsible for white fir and yellow pine invasion of sagebrush-dominated associations at lower-elevation sites in the Warner range. Pure whitebark stands appear to be self-maintaining, as seedling density beneath mature individuals is high and consists solely of whitebark pine seedlings throughout the pure stands in the study area.

Duffield, W.A. and R.D. Weldin. 1976. Mineral resources of the South Warner Wilderness, Modoc County, California. Calif. Geol. Surv. Bull 1385-D U.S. Govt. Printing Office, Washington, D.C.

Macdonald, G.A. and T.E. Gay, Jr. 1966. Geology of the southern Cascade range, Modac Plateau and the Great Basin areas in northeastern California. Pp 43-48 in Mineral resources report part I. Calif. Div. of Mines and Geol. Sacramento, CA.

Mattson, D.J. and D.P. Reinhart. 1990. Whitebark pine on the Mt. Washburn massif, Yellowstone National Park. In Proceedings-Symposium on Whitebark pine ecosystems: ecology and management of a high-mountain resource. U.S.D.A. Forest Service, Intermountain Research Station, General Technical Report INT-270.

Riegel, G.M., D.A. Thornburgh, and J.O. Sawyer. 1990. Forest habitat types of the south Warner Mountains, Modoc County, California. Madrono 37:88-112.

Vale, T.R. 1975. Invasion of big sagebrush by white fir on the southeastern slopes of the Warner Mountains, California. Great Basin Naturalist 35:319-324.

Vale, T.R. 1977. Forest changes in the Warner Mountains, California. Annals of the Association of American Geographers 67:28-45.

Whitebark Pine Health in Northern Rockies National Park Ecosystems: A Preliminary Report by Katherine Kendall, David Schirokauer, Erin Shanahan, Rob Watt, Dan Reinhart, Roy Renkin, Steve Cain and Gerry Green

In 1995, we began a three-year project to look at the status of whitebark pine in national parks in the Rocky Mountains from Wyoming to Alberta. During the 1995 and 1996 field seasons, we collected detailed information on tree status and the effect of damaging agents in 316 whitebark pine stands. Those data are reported here. Field work in 1997 will focus on; 1) validating the maps that we are creating this winter of current and historical whitebark pine distribution in Glacier NP and 2) repeating historical blister rust and ribes incidence surveys in Glacier and Yellowstone NP’s.

In general, whitebark pine trees in Waterton Lakes NP, Alberta, and Glacier NP and Blackfoot Reservation, Montana have suffered serious declines, while further south, whitebark pine mortality was low and blister rust infection rates were moderate to low (Table 1). In the northern portion of our sample area, approximately 30% of the whitebark pine trees were dead, and of the remaining live trees, about 70% were infected with rust and had an average of 25% crown kill.

In the Greater Yellowstone Ecosystem, whitebark mortality averaged 7%. Blister rust infection of whitebark pine occurs throughout the area, mostly at low levels (<5%). Infection rates in some areas were considerably higher. In Grand Teton NP, for example, blister rust incidence averaged 10% but infection rates in several sites ranged from 40-44%. The moister climate of the Tetons compared with Yellowstone NP is probably responsible for the higher rust levels.

The whitebark pine mortality and blister rust incidence levels reported here should be considered minimums. Seedlings and sapling whitebark pine trees which have been dead for a while often could not be identified to species or tended to be broken off at ground level and were not included in our whitebark pine mortality figures. Blister rust incidence reported in Table 1 as the "definite infection" rate include only those trees which were obviously infected with rust. Often, we encountered trees with evidence of rust, such as dead tops, flagged branches, and rodent gnawing of suspected cankers, but no definite cankers. When these "probable infections" were included with definitely infected trees, blister rust rates rose in all areas (Table 1).

Table 1. Preliminary data on the status of and effect of blister rust on whitebark pine (Pinus albicalus) stands sampled in 1995–96.

Area sampled	Number of sample sites	Mean Mortality %¹(range)	Mean Definite infectn % (range)	Mean Probable infection %²(range)	Mean crown kill %³(range)
Blackfoot Res. MT	5	61 (45-85)	84 (65-100)	85 (69-100)	30 (18-52)
Central Idaho	1	1	3	5	0
Glacier NP, MT	174	45 (0-100)	73 (0-100)	79 (0-100)	29 (0-70)
Grand Teton NP, WY	34	7 (0-50)	10 (0-44)	15 (0-62)	1 (0-9)
Waterton L. NP, Alberta	8	26 (14-48)	44 (14-59)	47 (14-62)	13 (3-18)
Yellowstone NP, WY, MT, ID	95	7 (0-64)	4 (0-58)	5 (0-58)	1 (0-21)
¹ Includes mortality from all causes. ²Includes trees which were probably, as well as, definitely infected with blister rust. ³ Average crown kill of all live whitebark pine trees.

Whitebark Pine Disease Survey for the Northern Portion of The U. S. Forest Service Intermountain Region by Jonathan Smith and Jim Hoffman, University of Idaho, Dept. Forest Resources, Moscow, ID 83843, Phone: 208-883-8686, Email: smit9423@uidaho.edu

Recent concern over whitebark pine (Pinus albicaulis) decline prompted the U.S. Forest Service’s Forest Pest Management Office to take a closer look at the whitebark pine in their region. The Intermountain region, Region 4, encompasses the national forests of Idaho south of the Salmon River, those of Nevada, Utah, and western Wyoming. We spent the past two summers surveying whitebark pine stands for disease incidence throughout the range of whitebark pine within this region.

The goal of the survey is to investigate the distribution of diseases, especially blister rust disease (caused by the introduced fungus Cronartium ribicola), in whitebark pine forests of the region. In the absence of baseline data, it is difficult for forest managers and researchers to identify where blister rust disease is currently affecting this important ecosystem. Our survey will describe the distribution of blister rust disease and attempt to determine the current status of the disease within the region.

In addition to describing the distribution pattern, we are investigating potential relationships between the distribution of blister rust disease and site and stand factors. If these factors are related to incidence and severity, data such as ours could help land managers make decisions about where to focus restoration efforts.

We randomly selected sample stands that were identified by local land managers as reasonably accessible. We then systematically located transects within each stand based on several sampling criteria. We collected data on the following variables:

The map shows the location of our 79 sample sites, and the approximate blister rust incidence level (percent of trees infected with rust) for each location. Additionally, we noted blister rust disease incidence in limber pine (Pinus flexilis) stands that we encountered en route to our sample stands. Limber pine stands are classified on the map as either infected or uninfected.

Concentrations of high blister rust disease incidence can be seen in western (Payette N.F.) and eastern (Targhee N.F.) portions of the study area. Disease incidence generally decreases as one travels southward in the study area, but blister rust disease was observed essentially throughout the range of whitebark pine in the region. Low incidence levels are concentrated in the mountain ranges of the Sawtooth and Challis National Forests in south-central Idaho.

Blister rust-related mortality or crown kill was nonexistent in most stands that we sampled, but some mortality is beginning to occur in the Centennial Mountains (Targhee N.F.) and to a lesser degree in the Bitterroots, Salmon River Mtns. and in the Gros Ventre and Wyoming Ranges (Bridger-Teton N. F.) This mortality is generally restricted to a few smaller diameter trees (less than 4 inches DBH) that have been girdled by blister rust cankers.

Overall, it appears that blister rust disease incidence is widespread and higher than we expected in most areas of the region. The very high incidence level in the Greater Yellowstone Ecosystem area (Targhee N.F.) is of special concern because of the substantial grizzly bear population in this ecosystem. Whitebark pine seeds are an important food source for these animals. In the coming months we will be investigating the distribution patterns as well as summarizing and analyzing the site and stand data. Questions and comments are welcome.

Jonathan P. Smith James T. Hoffman, Plant Pathologist
Department of Forest Resources U. S. Forest Service
University of Idaho Forest Pest Management
Moscow, Idaho 83844 Boise, Idaho 83702
208/885-7509 208/364-4221
J.Smith:R04F15D02A J.Hoffman:R04F02A
smit9423@uidaho.edu

Preliminary Status Report on Whitebark Pine in Gallatin National Forest, Montana by Katherine Kendall, Dan Tyers, and David Schirokauer, Biological Resources Division, USGS, Glacier National Park, West Glacier, MT 59936-0128, Phone: 406-888-5441
Email: katherine_kendall@usgs.gov

In 1995 and 1996, whitebark pine trees were surveyed in 47 stands in the Gallatin NF to determine status and degree of blister rust present. More than 90% of these sites were in the Gardiner District, adjacent to the northeast corner of Yellowstone NP. We collected data on site attributes and whitebark pine status, size class, crown ratio and class, and type and severity of damage as described in Kendall (1995). Volunteers, trained in field procedures and blister rust identification in whitebark pine, accomplished much of the field work. They were instructed to record a tree as infected with rust only when an unmistakable canker was present. Because of this conservative approach and their relative inexperience identifying rust, our estimates of infection rates are probably low.

Mean mortality of whitebark pine trees from all causes was 8%. The proportion of trees definitely infected with blister rust averaged 2% across all plots. These rates were comparable to those found in Yellowstone NP and were 1/2 - 1/10 of the infection rate found in the Tetons (Kendall et al., 1996).

Table 1. Preliminary results from the 1995-96 survey of the status and effects of blister rust on whitebark pine (Pinus albicalus) stands in the Gallatin National Forest.

Kendall, K.C. 1995. Whitebark pine monitoring network guide. Glacier Field Station, National Biological Service, Glacier National Park, MT 33p.

Kendall, Katherine, David Schirokauer, Erin Shanahan, Rob Watt, Dan Reinhart, Roy Renkin, Steve Cain and Gerry Green. 1996. Whitebark pine health in northern rockies national park ecosystems: A preliminary report. USDA Forest Service, Intermountain Research Station, Nutcracker Notes 7.

An Outstanding Prescribed Burn in Whitebark Pine at the Stevensville Ranger District, Bitterroot National Forest by Bob Keane and Steve Arno, Intermountain Fire Sciences Lab, P.O. Box 8089, Missoula, MT 59807, Phone: 406-329-4846

Little did she know it, but on October 2, 1996, Assistant Fire Management Officer Leslie Anderson ushered in a new era in whitebark pine management by lighting one of the first successful prescribed fires in a mature whitebark pine stand. Better yet, this stand actually contained two units side-by-side -- one a natural stand and the other a group selection cut to favor whitebark pine. The stand is about 10 acres in size at about 7200-7400 feet elevation on moderately steep south-facing slopes on the east slope of the Bitterroot Range. This is a mixed species, 200 year old stand of whitebark pine, lodgepole pine, subalpine fir and spruce. Undergrowth is primarily tall huckleberry, whortleberry, and beargrass. While the upper unit was a natural stand that was underburned, and the lower unit was logged in autumn of 1995 to create small openings to induce nutcracker caching. Trees were thinned between openings with all commercial fir and spruce and some lodgepole removed and whitebark pine was saved. Slash was left on the unit. Both units were broadcast burned.

Ignition began at top of upper unit at 12:55 pm and continued slowly until 7:30 pm when the bottom edge was lit. Weather was partly sunny, warm and dry from late morning until dusk with thin high clouds and light winds. Temperatures ranged from 51 degrees at the start to a maximum of 60 degrees, while relative humidies ranged from 31 to 21 percent. Weather had been warm dry and often windy for several days, with 0.96 inches of precipitation during the previous 3 weeks as measured at a remote (RAWS) station immediately below the burn site. Log moisture contents ranged from 16 to 28 percent while fine woody fuel (< 1 inch dia) moisture contents ranged from 14 to 33 percent. Lighting began with drip torches at upper east corner and almost immediately, a tall, old subalpine fir torched out shooting flames up over 100 feet into the air. The shower of burning embers landed in various fuels across the fire line but only rarely did these embers ignite a spot fire. Burning the upper unit was done slowly with narrow strips to keep fire intensities low. Old firs often torched out spectacularly, but small firs seldom burned apparently not having sufficient dry fuel accumulations around the base. Huckleberry leaves had been killed by a hard frost in September and supported sporadic low-intensity burning as did the thin dry duff layer. Old down tree trunks often burned vigorously. Areas having any concentration of dead material and old firs burned with moderate intensity; other areas burned minimally in the natural stand.

The logged unit had light to moderate loadings of cured slash (about 20 tons per acre) and this burned vigorously covering most of the area. Fire failed to spread into the small riparian strip in the center of the logged unit; there was some flowing seep water here and no logging or slash. Despite occasional showers of embers from torching fir and spruce that landed in the control unit east of the fire line only a few slow-developing spot fires happened and they were easily controlled.

The fire continued to smolder for the next four days, and additonal burnout of ground fuels and torching of subalpine firs occurred with afternoon heating. It was estimated that over 50 percent of the surface in the natural stand unit experienced burning by two days after the burn. Burn coverage in the logged stand was estimated at about 90 percent. We have excellent video footage of the fire as well as numerous 35 mm. color slides. We will be monitoring the effects of this fire over the next ten years by measuring changes in tree growth, understory species composition and coverage, and fuel loadings. If fact, we observed nutcracker caching in the logged unit nutcracker openings during the two weeks prior to the burn.

Leslie Anderson, Bitterroot NF Silviculturalist Cathy Stewart, and the staff at Stevensville Ranger Station did a great job planning and implementing this important prescribed burn which, in addition to being an important research site, will be a focal point for future field demostrations because of its accessibility. By all accounts this fire met and exceeded all burn objectives, and hopefully, its success will confirm the use of prescribed fire as an important tool for whitebark pine ecosystem restoration.

INT Cone Collection Update: The seed year we’ve all been waiting for! by Donna Dekker, Forestry Sciences Laboratory, Moscow, ID 83843, Phone: 208-883-2324, Email: d.dekker:s22l04a

For once the gods of Budget, Personnel and Cone Crop coincided. It was an excellent cone year over much of the region and the Intermountain Station made the most of it. In Nutcracker Notes #6 (December 1995), I wrote about the ongoing effort to collect whitebark pine seed across Region 1 for an adaptive variation study. Scientists at the Intermountain Station have been working on a region-wide collection of whitebark pine seeds since the early 1990's. At each collection site (typically a ridgetop or drainage) 10-11 conebearing whitebark pines are selected and the open-pollinated seed is collected. This seed will be planted in a common garden and measured for certain phenological traits such as date of budbreak, date of budset, and shoot elongation time. These, in turn, should give us some data from which to develop seed transfer guidelines.

The field season got a slow start in some areas because of last winter’s heavy snowpack. Flood damage closed roads across the region. The Intermountain team worked from north to south and east to west to leave the areas most heavily hit by winter weather (particularly the Nez Perce and Clearwater NFs) time to clear. Collections by the Intermountain team were made on the Bonner’s Ferry and Sandpoint districts of the Idaho Panhandle NF, the Missoula, Ninemile, Plains/Thompson Falls and Superior districts of the Lolo NF, the Stevensville, Sula, and West Fork districts of the Bitterroot NF, the Red River, Elk City, and Salmon River districts of the Nez Perce NF, and the Powell district of the Clearwater NF. Many thanks are extended to Kelly Smith (Bonner’s Ferry), Brian Dreisbach (Plains/Thompson Falls), Wes Paulson and Karen Harvey (Powell) and Joe Hughes (Superior) for their help in the field. Many others were extremely helpful when we stopped in the local district offices to locate the best stands on their districts, giving good advice, detailed maps, and permits for closed roads.

On the west side of the Continental Divide, the cones went to those who caged. Hardware cloth cone cages were used to protect the cones on most sites, but the warm, dry summer weather had the nutcrackers pecking at the cones by the end of July. Even though trees in many areas were loaded with cones, the nutcrackers and red squirrels didn’t leave any behind for collectors on most sites. By the time the teams arrived to remove the cone cages every unprotected cone was gone. On the east side of the Divide, rifle collections were successful although the cones were slow to ripen.

In general, climbable trees with cone crops were found on southern aspects near the summit. Trees on the northern aspects were often embedded in forests of subalpine fir, and were usually too tall to be easily climbed. When road access was available it worked best to drive to the top, then work our way down, caging cones as we went.

Additional collections were made by NFS personnel from several forests. The Gallatin NF’s Lars Halstrom was the busiest; his team collected cones from seven sites east of the Divide. The Gallatin team made collections on the Bozeman district, on the Wise River and Madison districts of the Beaverhead NF, on the Island Park district of the Targhee NF, and on the Lincoln district of the Helena NF. They also traveled to the Sweetgrass Hills, a barren outcrop just south of the Canadian border and well east of Glacier NP.

The Clearwater NF’s Jerry Branning (North Fork district) made a collection on Blacklead Mountain, while the Powell district’s Wes Paulson, Karen Harvey and John Weston made a collection at Beaver Ridge Lookout. Intermountain Fire Sciences Lab scientist (and Nutcracker Notes editor) Bob Keane got a collection in the Sapphire Range on the Bitterroot NF.

We were skunked on a few sites west of the Divide, but nowhere more than on the Flathead and the Kootenai national forests. On a few sites a lone tree with cones was found, but it was clear that we missed the boat by not making more collections last year. Hopefully it will be possible to pick up a few sites in the summer of 1997.

All in all, we now have more than 70 collections which, provided that the seed yield this year is good, is enough to sow the adaptive variation test. I would like to be able to add another collection or two from northwestern Montana so that part of the region is better represented. However, the desire for more collections must be balanced by the decreasing viability of seed in storage. Since 1996 was such a good cone year through much of the region, and 1995 was good in northwestern Montana, the odds are against another good cone crop for a couple of years.

Editors Note: Indeed, this year was a very good year for whitebark pine cones. But how "good" was it? It seems "highly variable" might be the best term to describe this year. Comments about the 1996 whitebark pine cone crop were solicited from people on the whitebark pine electronic mailing list and the following is a summary of the survey results. In Montana, whitebark pine stands west of the Continental Divide had good to excellent cone crops. Good cone crops were observed in the Bitterroot, Sapphire, Swan, and Mission Ranges. Steve Arno mentions that the Bitterroot Front cone crop was much heavier north of Hamilton. Kate Kendall found a poor crop in Glacier National Park, and many people observed moderate to good crops in northern Idaho. However, cone crops east of the Divide were incredibly variable. Lars Halstrom notes that cone crops in the Centennial, Gallatin and Madison Ranges were poor to moderate, with small and under-developed cones that had many empty seeds. Jon Smith found large to massive cone crops on the Bridger-Teton National Forest, moderate crops on the Targhee National Forest, and poor to good crops on the Salmon-Challis National Forests. The Boise National Forest had only light cone crops observed from a limited number of sites. Barbara Levesque and fellow workers found near bumper cone crops on the Cobalt District of the Salmon-Challis NF. I observed moderate cone crops in the northern and central Selway-Bitterroot Wilderness Area. Melissa Jenkins notes a very good cone crop on the Island Park District of the Targhee NF. Brian Kelley and Bud Kovalchik observed a very good cone crop in the Lake-Chelan-Sawtooth Wilderness on the Okanogan NF.

Limber Pine Status from Alberta to Wyoming by Katherine Kendall, Dayna Ayers, and David Schirokauer, Biological Resources Division, USGS, Glacier National Park, West Glacier, MT 59936-0128, Phone: 406-888-7994, Email: katherine_kendall@usgs.gov

Limber pine is a 5-needled pine widely distributed in the mountains and foothills of the western United States and southern Canadian Rocky Mountains. Little is known about limber pine ecology and condition. During our studies of whitebark pine in Glacier National Park, MT, we became concerned about the high amount of mortality in limber pine stands. Like its cousin whitebark pine, it is highly susceptible to white pine blister rust. In 1995 and 1996, we recorded site characteristics, tree status and damaging agents in 81 limber pine stands from southern Alberta to eastern Idaho and northern Wyoming. Our sampling was far from comprehensive but we believe the results reflect the general pattern of limber pine tree status in this region.

In the areas we sampled, limber pine occurred on extremely xeric sites at lower and upper tree line. We usually found lower treeline stands in areas too dry for ponderosa pine, such as the east slopes of the Beartooth Plateau and various island mountain ranges. Pure stands of limber pine were commonly located on windswept, rocky ridges, whereas mixed stands were more common on hillsides, usually with south-facing slopes. Occasionally, mixed stands of limber pine and ponderosa pine, Douglas fir, or lodgepole pine were located on slopes with northern aspects where soil moisture was higher. Whitebark pine was mixed with limber pine in many stands sampled on the Blackfoot Indian Reservation and east side of Glacier NP in Montana.

Limber pine has suffered extensive, heavy mortality and blister rust infection in northwest Montana and southern Alberta (Table 1). On average, over a third of the limber pine is dead and about 75% of the remaining live trees are infected with rust and 30% of the crown has been lost. Limber pine health improves somewhat to the north and south. Mortality and blister rust incidence rates were lower north of Waterton Lakes in the Porcupine Hills, Alberta. In southwest Montana, northwest Wyoming, and adjoining areas of Idaho, limber pine mortality and incidence of rust is low to moderate with a few hot spots of heavy infection. Although blister rust incidence in limber pine stands in the Bighorn Mountains of north-central Wyoming is generally low, we found high infection rates and significant mortality at sites in the northeast and southwest corners. We found no rust in Craters of the Moon National Monument in southern Idaho. When trees identified as "probably infected" were included, rust infection rates rose for all areas, sometimes substantially, as in the Porcupine Hills (Table 1).

Not all damage to limber pine trees was attributed to blister rust. In some stands with dead and dying trees or trees with thinning crowns or dead tops, we saw no definite cankers or other evidence of rust. Some of the defoliation was due to limber pine needle cast but a number of factors, such as severe climatic events, mistletoe, and/or mountain pine beetles, may have combined to cause the poor health observed (McConnell 1996).

Whitebark pine seeds are a well documented bear food but reports of bears feeding on limber pine seeds are scarce. We found evidence that bears ate limber pine seeds in the Porcupine Hills (2 scats) and in Waterton Lakes NP, Alberta (1 scat). The scats were found in limber pine stands with no whitebark pine trees in the vicinity. Many limber pine trees we surveyed were short enough that bears could reach cones without climbing trees. Thus, limber cones were available to grizzly bears (poor climbers) without squirrels acting as intermediaries by cutting down cones.

McConnell, T. 1996. Montana forest insect and disease conditions and program highlights -1995.USDA, Forest Service, Forest Health Protection, Report 96-2:19-20.

Kate Kendall, Research Ecologist,
Biological Resources Division, USGS,
Glacier National Park,
West Glacier, MT 59936-0128.
katherine_kendall@usgs.gov

David Schirokauer, Biologist,
Biological Resources Division, USGS,
Glacier National Park,
West Glacier, MT 59936-0128.
dave_schirokauer@usgs.gov

Dayna Ayers, Boone and Crockett Wildlife Conservation Program,
School of Forestry, University of Montana,
Missoula, MT 59812.
dayna@selway.umt.edu

Hutchins, Harry E., S.A. Hutchins, B. Liu. 1996. The role of birds and mammals in Korean Pine (Pinus koraiensis) regeneration dynamics. Oecologia 107:120-130.

Lanner, Ronald M. 1996. Made for each other -- a symbiosis of birds and pines. Oxford University Press. 160 pages.

Mattson, David J., and Daniel P. Reinhart. 1996. Indicators of Red Squirrel (Tamiasciurus hudsonicus) abundance in the whitebark pine zone. Great Basin Naturalist 56(3):272-275.

Murray, Michael P. 1996. Landscape dynamics of an island range: Interrelationships of fire and whitebark pine (Pinus albicaulis). Doctoral Dissertation. University of Idaho, Moscow, ID.

Krebill, R.G. and R.J. Hoff. 1995. Update on Cronartium ribicola in Pinus albicaulis in Rocky Mountains, USA. Proc. 4th IUFRO Rusts of Pines Working Party Conf., Tsukuba: 119-126.

Tomback, D.F., K.S. Carsey, and M.L. Powell. 1996. Post-fire patterns of whitebark pine (Pinus albicaulis) germination and survivorship in the greater Yellowstone area. In: J. Greenlee (Editor), Proceedings of the second biennial conference on the Greater Yellowstone Ecosystem, Yellowstone National Park, Wyoming, Sept. 19-21, 1993. International Association of Wildland Fire, Fairfield, WA, USA. Pages 21-31.

The symposium: "Restoring Whitebark Pine Ecosystems -- A Field Workshop" is now scheduled for September of 1998 because research project results had been delayed and because of budget constraints. The tentative dates for this symposium are Sept 8-11, 1998. More information about this symposium will be published in the following issues of Nutcracker Notes.

Interesting Talk: Ron Lanner, Professor Emeritus of Forest Resources at Utah State University, will be in Missoula on March 11, 1997 to present a talk about the symbiotic relationship of Corvids and Pines which is the subject of his new book "Made for each other -- a symbiosis of birds and pines" (see above). This will be an evening keynote speech given at the Bitterroot Ecosystem Management Research Project annual conference. This date is tentative and subject to change. Contact Bob Keane or Steve Arno for details (406-329-4800).

Bob Bell, a graduate student at UCLA currently working in Yellowstone National Park, is constructing a home page for whitebark pine on the World Wide Web. Once finished, he will pass the page over to the Forest Service and they will maintain it on their home page. All editions of Nutcracker Notes will be available at this site, along with many other interesting things about this ecosystem. Got an ideas? Get them to Bob Keane and he will try to add them to the web page.

NUTCRACKER NOTES is a vehicle for the dispersal of information on all facets of whitebark pine ecosystems. Summaries of research results and management projects in whitebark pine forests are presented to provide readers state-of-the-art information. The purpose of this newsletter is to distribute timely information so that land managers and scientists can understand and deal with important ecological issues in the whitebark pine ecosystem. Issues of NUTCRACKER NOTES will be numbered and published 1-3 times a year depending on available material.

Submission of Articles: Everyone is invited to submit articles to NUTCRACKER NOTES. These articles should be mailed to Nutcracker Notes, c/o Bob Keane, Intermountain Fire Sciences Lab, P.O. Box 8089, Missoula, MT 59807. If possible, they should be submitted electronically to B.KEANE:S22L01A over the Data General, or written to a floppy disc (WordPerfect text processing) and then mailed. You are encouraged to submit articles to improve this information network.

Newsletter Format: Articles submitted to NUTCRACKER NOTES will be presented in the newsletter under three main categories: Management News and Notes, Research News and Notes, and Publication and Events Alert. Management News describes current activities, problems, observations, conditions planned or implemented by land management agencies in whitebark pine forests. Research News describes current or planned research projects in these ecosystems. Publication and Events Alert is simply a list of current events and published information that may be of interest to readers of the newsletter. At the end of the newsletter the reader will find a complete list of all authors along with their addresses. There will usually be an editorial at the beginning of the newsletter to highlight important topics and provide a forum for opinions.

Errata and omissions: Issue #6 -- Dr. Donna Dekker’s address was omitted from the previous issue. It can be taken from the Editorial she has written in this issue number seven..

Table 1. Whitebark pine mortality data for six eastern Washington cascade sites
Sites	Location	No. Trees	Alive	Recent Dead	Old Dead	Very Old Dead
Mt Adams	Yakama Ind. Res.	51	47	1	1	2
Goat Butte	Yakama Ind. Res.	50	47	1	1	1
Trinity Mtn	Okanogan NF	104	102	1	1	0
Mission Ridge	Wennatchee NF	120	103	2	5	10
Stormy Mtn	Wenatchee NF	59	52	1	4	2
Shady Pass	Wenatchee NF	78	72	1	2	3
Total		462	423	7	14	18

Table 2. White pine blister rust infection in different whitebark pine size classes in six eastern Washington Cascade sites.
Site	Trees*	Seedlings	Saplings	Poles	Mature
Mt Adams	51-10	20-0	13-6	7-4	11-0
Goat Butte	50-13	11-0	26-8	6-3	7-2
Trinity Mtn	104-28	22-2	31-5	24-9	27-12
Mission Ridge	120-16	22-1	70-8	15-4	13-3
Stormy Mtn.	59-11	12-1	23-6	5-1	19-3
Shady Pass	78-15	17-0	36-10	10-1	15-4
Totals	462-93	104-4	199-43	67-27	92-24
* First number is the total trees, the second is the number infected by blister rust.

Table 1. Preliminary data on the status of and effect of blister rust on limber pine (Pinus flexilus) stands sampled in 1995–96.
Areas sampled	Number of sample sites	Mean Mortality %¹(range)	Mean Definite infection % (range)	Probable infectn %²(range)	Mean crown kill %³ (range)
Bighorn Mtns., WY		2 (0–6)	22 (0–58)	32 (4–68)	4 (0–15)
Blackfoot Reservation, MT	6	42 (0–68)	53 (0–100)	53 (0–100)	27 (0–56)
Central Idaho	2	0	0	0	0
E. Front, MT	3	21 (16–24)	14 (4–27)	30 (23–43)	8 (7–9)
N.W. Wyoming	13	10 (0–62)	24 (0–78)	28 (0–87)	5 (0–19)
Porcupine Hills, Alberta	4	14 (4–23)	48 (43–57)	76 (64–88)	22 (12–28)
Grand Teton NP, WY	1	16	32	32	6
Glacier NP, MT	22	39 (0-89)	78 (0-100)	80 (0-100)	35 (0-80)
Yellowstone NP, WY, MT, ID	3	28 (4-41)	1 (0-2)	2 (0-6)	0 (0-0)
Gallatin NF, MT	4	37 (9-88)	17 (0-39)	24 (2-46)	15 (13-17)
S.W. Montana	11	19 (2–88)	29 (0–89)	33 (0–89)	7 (0–19)
Waterton L. NP, Alberta	8	46 (26–85)	85 (65–97)	93 (81–98)	33 (19–49)
¹Includes mortality from all causes. ² Includes trees which were probably, as well as, definitely infected with blister rust. ³ Average crown kill of all live limber pine trees.