United States

Department

of Agriculture

Forest Service

Rocky Mountain

Research Station

General Technical

Report RMRS-GTR-208

April 2008

Death of an Ecosystem:

Perspectives on Western White Pine 

Ecosystems of North America at the 

End of the Twentieth Century

Alan E. Harvey, James W. Byler, Geral I. McDonald, 

Leon F. Neuenschwander, Jonalea R. Tonn
Rocky Mountain Research Station

Natural Resources Research Center

2150 Centre Avenue, Building A

Fort Collins, Colorado 80526

Harvey, Alan E.; Byler, James W.; McDonald, Geral I.; Neuenschwander, Leon F.; 

Tonn, Jonalea R. 2008. Death of an ecosystem: perspectives on western white 

pine ecosystems of North America at the end of the twentieth century. Gen. 

Tech. Rep. RMRS-GTR-208. Fort Collins, CO: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station. 10 p.

The effective loss of western white pine (Pinus monticola Dougl.) in the white pine 

ecosystem has far-reaching effects on the sustainability of local forests and both 

regional and global forestry issues. Continuing trends in management of this forest 

type has the potential to put western white pine, as well as the ecosystem it once 

dominated, at very high risk in the future. Societal issues associated with natural 

resource management must be resolved early in the 21st century to allow restoration 

of this ecosystem so that the Interior Northwest’s most productive forests can be 

sustainable at levels near their historical potential.

Keywords: white pine blister rust, ecosystems, Ribes, western white pine

Alan E. Harvey is Supervisory Plant Pathologist, retired, U.S. Department of 

Agriculture, Rocky Mountain Research Station, Forestry Sciences Lab, Moscow, ID.

James W. Byler is Group Leader, retired, U.S. Department of Agriculture, Forest 

Service, Northern Region, Forest Pest Management Field Office, Coeur d’Alene, ID.

Geral I. McDonald is Principal Plant Pathologist, emeritus, with the U.S. Department 

of Agriculture, Forest Service, Rocky Mountain Research Station, Forestry Sciences 

Lab, Moscow, ID.

Leon F. Neuenschwander is Professor, retired, of Forest Resources, University of 

Idaho, Moscow, ID.

Jonalea R. Tonn is Forester with the U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Forestry Sciences Lab, Moscow, ID.

Abstract

The Authors

The authors are grateful for comments contributed by Dr. Dennis Ferguson, Dr. 

Russell Graham, Dr. Raymond Hoff, and Dr. Ned Klopfenstein. Thanks go to John 

Hanna for assisting with formatting the photographs. Funding for this work was 

provided by the USDA Forest Service, Rocky Mountain Research Station, Forest 

and Woodland Ecosystem Program, Moscow, ID. Use of trade names does not 

constitute endorsement by the USDA Forest Service.

Acknowledgments

You may order additional copies of this publication by sending your 

mailing information in label form through one of the following media. 

Please specify the publication title and series number.

Fort Collins Service Center

Telephone (970) 498-1392

FAX (970) 498-1122

E-mail rschneider@fs.fed.us

Web site http://www.fs.fed.us/rm/publications

Mailing address Publications Distribution

Rocky Mountain Research Station

240 West Prospect Road

Fort Collins, CO 80526
Contents

Introduction............................................................................................................ 1

Western White Pine: the Species .......................................................................... 1

Historic Dominance of Western White Pine........................................................... 2

Nature of Western White Pine Dominated Ecosystems ........................................ 2

Introduction of White Pine Blister Rust.................................................................. 4

Current Status of Western White Pine................................................................... 6

Comparative Nature of Replacement Forests ....................................................... 7

Management Implications...................................................................................... 8

Perspectives About the Future of Western White Pine.......................................... 9

References ............................................................................................................ 9
Figure 1—Continuous forest cover of nearly pure western white pine in northern Idaho, 1938 (USFS file photo).
USDA Forest Service RMRS-GTR-208. 2008. 1

Introduction

Early in the century, western white pine (WWP) 

was a dominant, highly valuable, and easily cultured 

species that spawned forest industries in the West and 

represented a primary economic strength for the region 

(Neuenschwander and others 1999) (fig. 1). It was the 

key to imparting many extremely important characteristics to Interior Northwest forests (Harvey and others 

1995; Monnig and Byler 1992). Most important was 

its ability to form a stable, relatively long-lived, mixed 

species forest that was perpetuated by a combination 

of mixed-severity and stand-replacing wildfires (Zack 

and Morgan 1994). Even though fire often occurred 

in this forest type (25 to 30 yrs for mixed-severity 

and 150 to 200 yrs for stand-replacing fires), oldgrowth structures often persisted for several centuries. 

Although WWP remains potentially the most economically and certainly the most ecologically valuable 

species in much of the region, white pine blister rust 

(WPBR) has reduced it to a relic. Consequently, WWP 

is now largely represented as widely scattered trees 

with limited natural regeneration potential in areas it 

formerly dominated (Atkins and others 1999; Harvey 

and others 1995; Monnig and Byler 1992).

Western white pine will be difficult to re-establish 

without strong political will. In the absence of aggressive culture, it will shortly be lost as a significant 

ecosystem component throughout much of its range 

(Neuenschwander and others 1999). In its absence, 

forest ecosystems of the region will be much less productive. These ecosystems may only be sustainable 

as low productivity, unstable, 

and insect-pathogen-fire-prone 

forests of relatively low value 

for all amenities, including watershed protection and carbon 

sequestration (Atkins and others 1999; Harvey and others 

1995; Monnig and Byler 1992).

Western White Pine: 

the Species

Western white pine is the 

key to a uniquely valuable, productive, and stable forest type 

that was once prominent in the 

Interior Northwest (fig. 2). In 

northern Idaho and contiguous portions of Washington, 

Montana, and British Columbia, 

the species often dominated most of the highly productive sites. It always functions as a seral component. 

It is a tall, straight tree that self prunes well with relatively open, low-density foliage (Graham 1990). 

When less than mature, it is highly tolerant of most 

endemic insects and pathogens, especially when compared with late seral and climax species that are its 

most common associates (Harvey and others 1995).

Where WWP has adequate light, it is a good seed 

producer (fig. 3). The cones are large, up to 33 cm (13 

inches) long, and well filled with seed. The large seed 

germinates well in the spring. Germination and early 

growth are especially aggressive on burned surfaces. 

The seed is unique to the region in that portions can 

remain viable in the soil more than 1 year. Seedlings 

survive best with substantial sunlight, are deep rooted, 

and survive drought moderately well when compared 

to most of their common associates. As a mature tree, 

it is only moderately shade tolerant and does not 

produce seed well when not achieving at least a codominant position (Haig and others 1941; Miller and 

others 1927).

The species is effectively perpetuated only through 

stand-opening disturbances. Historically, this occurred through fire or insect and pathogen activities, 

mostly the latter, as they affected competing species. 

More recently, harvesting and subsequent management disturbances have been effective at propagating 

the species (Graham 1990; Neuenschwander and others 1999).

Western white pine was, and still can be, the most 

valuable commercial species in the region. It provides 

a light, strong, and straightgrained wood product that 

holds paint well, does not 

easily split, and is particularly amenable to production 

of high-value sash, paneling, and molding, as well as 

products such as matches and 

tooth picks. In the early part 

of the century (prior to the actions of WPBR), it was highly 

amenable to management, responding well to clear-cuts, 

seed-tree cuts, open shelterwood cuts, or similar systems 

popular at the time (Haig and 

others 1941; Miller and others 1927). It tends to be most 

prevalent on easily accessible, 

gentle slopes and on productive soils. For these reasons, 

Figure 2—Interior of an old-growth western white 

pine stand, 1935 (USFS file photo).
2 USDA Forest Service RMRS-GTR-208. 2008.

it has received considerable commercial attention and 

management in the region since the late 1800s (Graham 

1990; Haig and others 1941; Neuenschwander and 

others 1999).

Historic Dominance of 

Western White Pine

A century ago, WWP was a significant component 

of, or more often dominated, most of the productive, 

high-resource (moist, fertile) sites in the heart of the 

Inland Northwest. Before the 1860s, it made up 25 

to 50 percent of over 2 million ha (approximately 5 

million ac) of forest lands in the region. It varied in 

composition from 15 to 80 percent of the stands, sometimes even more, especially in northern Idaho. Where 

the composition of WWP was more than 15 percent of 

total basal area or 15 percent of total stems in regenerating forests, it was considered to be “white pine type” 

because of the economic importance and high interest 

in managing the species throughout the region (Miller 

and others 1927).

Nature of Western White Pine 

Dominated Ecosystems

Over the long term, WWP grows well compared to 

most associated species, especially in northern Idaho. 

Figure 3—Western white pine was an 

aggressive regenerator of clearcut on 

burned sites, 1910 (USFS file photo).
USDA Forest Service RMRS-GTR-208. 2008. 3

Its growth rate in the heart of its range is challenged 

only by western larch (Larix occidentalis Nutt.), but 

in a limited portion of that range. The tree is tall, often 

reaching over 60 m (200 ft) (fig. 4). It can easily grow 

over 1 m (39 inches) in a year. At 30 years, it can be 

38 cm (l5 inches) in diameter and 20 m (65 ft) tall. At 

300 years, it can be 152 cm (60 inches) in diameter, 

60 m (200 ft) tall, and still be gaining height. It often 

produces 48 m3

/ha (50,000 bd ft/ac) in 100 years. Some 

old-growth stands produce over 95 m3

/ha (100,000 bd 

ft/ac) (Miller and others 1927).

In mature stands, not only is WWP tall, but it is also 

deep rooted (Minore 1979). Thus, its physical architecture is such that it spreads its resources from deep in 

the soil to high in the air. Both are desirable characteristics for a species that is regenerated primarily by fire 

and competes best on highly variable, high-resource 

sites. Additionally, its genetic architecture allows it to 

be highly tolerant of site and environmental variation, 

both typical throughout its range (Rehfeldt 1994).

One of the most striking characteristics of WWP is 

its capability to thrive in a wide variety of sites and 

environments. Throughout its range in the West, it 

has been previously divided into only two seed zones. 

Other seral species of the region (western larch and 

ponderosa pine [Pinus ponderosa Douglas ex Lawson 

and C. Lawson]) have moderately wide seed zones, but 

even they are nowhere near as ecologically “flexible” 

Figure 4—Western white pine is tall, with a 

well dispersed, narrow canopy that self 

prunes well (USFS file photo).
4 USDA Forest Service RMRS-GTR-208. 2008.

as WWP. In contrast, most late seral and climax species are not as flexible with respect to “home” habitat 

(Rehfeldt 1994).

Native insects and pathogens are powerful background forces in WWP-dominated ecosystems. They 

tend to remove the late seral and climax species, such 

as Douglas-fir (Pseudotsuga menziesii [Mirb] Franco), 

grand fir (Abies grandis Dougl. ex. D. Don.), and 

western hemlock (Tsuga heterophylla [Raf.] Sarg.), 

from stands as they age. That process perpetuates 

WWP dominance, even at advanced ages, where other 

species might otherwise have an advantage (Harvey 

and others 1995). This creates a situation where the 

combination of disturbance-related initiation (primarily fire) and continuous pressure on other species by 

endemic biological forces generated a seral forest that 

was the most stable and productive forest type in the 

region. In the absence of a significant component of 

WWP or other seral species, current forests break up 

early and exhibit generally low productivity as a result 

of their incapacity to tolerate site/climate variation and 

susceptibility to endemic biological agents (Harvey 

and others 1995; Monnig and Byler 1992).

A strong tendency to be tolerant of endemic insects 

and pathogens is a characteristic generally typical of 

seral species in white pine country. This is particularly 

true of WWP. The species is usually quite tolerant of 

the myriad of foliar insects and root-rotting pathogens 

typical of the region. It is relatively susceptible to 

Armillaria root disease only when young (less than 25 

years) and to mountain pine beetle largely at advanced 

ages (over 140 years). Historically, mountain pine 

beetle was the primary agent responsible for breakup 

of over-mature (200 to 250 years old) WWP stands 

(Miller and others 1927).

In a very real sense, WWP appears optimally 

equipped to deal with a native range that is highly diverse with respect to site, soil, and climate and was 

continuously disturbed by fire, wind, and endemic 

biological agents. It was positioned to dominate high 

resource forests in the Interior Northwest and did 

so, even in the face of aggressive harvesting, until 

the introduction of WPBR and imposition of strict 

fire controls early in this century (Arno 1980; Maloy 

1997; Neuenschwander and others 1999).

Introduction of White Pine 

Blister Rust

In the early 1900s, forest pathologists were highly 

aware of the WPBR threat to both eastern and western white pine (Spaulding 1911). Despite a dedicated 

attempt to keep it out of North America, WPBR was 

found on cultivated currants (alternate hosts in the 

genus Ribes) in eastern North America by 1906 and 

in western North America in 1921 (Mielke 1943). 

Unfortunately, by the time it was discovered in the 

West in the vicinity of Vancouver, British Columbia, it 

had already been in the area for 10 years. At the time 

of its discovery in the West, WPBR had already spread 

into domestic and wild populations of currants (Ribes

spp.), several exotic white pines in the area, and native 

WWP (fig. 5). Despite sometimes heroic efforts to stop 

or eradicate WPBR on the domestic and wild currants 

that carried it, it rapidly spread southward and eastward to prime western white pine and sugar pine (P.

lambertiana Dougl.) country (Mielke 1943). It reached 

Oregon and Idaho in the 1920s, Glacier Park by 1939, 

and the southern Sierras by 1961 (Maloy 1997). In 

the 1990s, it continued to move into the Southwest on 

high-altitude, five-needled pines, especially whitebark 

Figure 5—White pine blister rust seedling infection showing 

aeciospore production.
USDA Forest Service RMRS-GTR-208. 2008. 5

pine (P. albicaulis Engelm.), limber pine (P. flexilis

James), and southwestern white pine (P. strobiformis Engelm.). Bristlecone (P. aristata Engelm.) and 

foxtail pines (P. balfouriana Grev. & Balf.) are also 

vulnerable (McDonald and Hoff 2000). It will likely 

place these resources at highly increased risk early in 

the 21st century.

Contrary to results achieved in eastern North 

America, control efforts centered on eliminating the wild currant hosts in the West, but they 

were not sufficiently effective to stop the rust and 

the effort was abandoned in 1968. Similarly, despite some early encouragement, treatment with 

the systemic antibiotics Actidione® and Phytoactin®

(registered trademark products of UpJohn Corp. 

for cycloheximide derivatives and Pabst Brewing 

Co. for polyamidohygrostreptin, respectively) were 

also found to be ineffective and were abandoned 

(Ketcham and others 1968; Maloy 1997). Potential 

alternatives for the latter approach are available but 

have not been widely used (Bérubé 1996; Harvey 

and Grasham 1979) (fig. 6).

Fortunately, early observation of resistance to 

WPBR in the native populations of WWP in the 

Interior Northwest proved correct. Beginning in the 

1950s, a systematic search for heritable resistance was 

undertaken and was successful. WPBR-resistant seed 

stocks derived from materials discovered in this effort became available in significant quantities in the 

1980s (Bingham 1983). These materials, and others 

Figure 6—Lethal white pine blister rust stem canker (photo by John W. Hanna, USFS).
6 USDA Forest Service RMRS-GTR-208. 2008.

discovered since, have become the linchpins of efforts 

to combat this destructive exotic disease (DeWald and 

Mahalovich 1997; Hagle and others 1989; McDonald 

and others 1991).

Current Status of Western White Pine

In effect, the introduction of WPBR completely 

changed the function of one of the most productive, 

stable, and tolerant (in terms of both management and 

natural disturbances) ecosystems in North America, if 

not the world. As a result, the ecosystem has changed 

to one dominated by other species. Current forests 

have quite different characteristics than those they replaced (fig. 7).

In terms of population size throughout its range 

(in the Interior Northwest), WWP is now estimated to 

be less than 5 percent of what it was at the turn of 

the 20th century (Neuenschwander and others 1999). 

Where risk of WPBR is extremely high, the current 

Figure 7—Old-growth western white 

pine stands had notably sparse 

understories with little fine fuel 

(USFS file photo).
USDA Forest Service RMRS-GTR-208. 2008. 7

population may be as little as 1 percent of what it once 

was (Hann and others 1997). Where risk of WPBR infection is low, the population may be as high as 10 

percent of that earlier in the 20th century.

Despite high growth potential, formerly WWPdominated forests are now one of the lowest storage 

compartments for fixed carbon in the northern Rocky 

Mountains (Birdsey 1992). Further, the first generation of resistant stock is performing to expectations 

only on low-risk sites. On high-risk sites, even resistant early stock types are suffering high mortality, 

though strikingly less than that of native stock types 

(McDonald and Dekker-Robertson 1998). Newer materials should fare better on high-risk sites (DeWald 

and Mahalovich 1997).

Comparative Nature of 

Replacement Forests

With the possible exception of western redcedardominated (Thuja plicata Donn) stands on especially 

moist sites, productivity, value, and stability of WWPdominated ecosystems exceeds that of other species 

combinations throughout the heart of its range. As a 

result of frequent actions of a variety of insects and 

pathogens, dominance by alternative species leads to 

significant losses in both productivity and longevity 

with associated accumulations of vegetative debris 

(Harvey and others 2000) (fig. 8). This change, in concert with the shallow rooting and low-density crowns 

Figure 8—Replacement forests now tend to have large volumes of fine “ladder” fuels (USFS file photo).
8 USDA Forest Service RMRS-GTR-208. 2008.

Figure 9—Collapse of an old-growth western white pine 

stand at Deception Creek Experimental Forest in northern 

Idaho, 1995.

of replacement (climax) species (Minore 1979), appears to lead to a more strongly horizonated soil with 

larger accumulations of litter on the surface than characteristic of forests dominated by seral species (Harvey 

and others 2000). These situations will likely lead to 

rapid immobilization of nutrients, especially nitrogen, 

in surface horizons. Located at the surface, it is then 

subject to loss associated with any severe disturbance, 

especially fire. In the absence of disturbance, higher 

water tables and nutrient tie-up can quickly lead to 

vegetative stagnation in moist, cool forests, perhaps 

with in a single generation (Bormann and others 1995; 

Kimmens 1994).

The potential, and perhaps ultimate outcome, is 

a forest dominated by species with high nutrient demands where nutrient storage and cycling rates are 

increasingly depressed. This will likely lead to everincreasing stress, with associated endemic insect 

and pathogen activities creating a domino effect that 

destabilizes the ecosystem (for example, excessive 

mortality and more frequent fire). The destabilized 

ecosystems exhibit inappropriate sensitivity and longterm damage from the same disturbances that once 

created a highly productive and stable forest ecosystem that was well adapted to both the characteristic 

long fire cycles and the activities of native insects and 

pathogens.

As WWP increasingly loses its ability to attain a 

codominant position, it will lose its ability to produce seed. Without substantial openings, any of the 

shade intolerant pine seedlings that are produced will 

quickly lose out to competition from large numbers 

(4 to16,000 ha [10 to 40,000 ac]) of shade-tolerant fir 

and hemlock (Graham 1990). Therefore, even without 

the ravages of WPBR, low numbers of scattered WWP 

may not be sustainable in these forests over the long 

term.

Perhaps most important in this species conversion 

process is the resulting change in genetic strategy 

of the primary vegetative components. This change, 

from one of wide adaptability and tolerance for endemic insects and pathogens to one of narrow adaptive 

capacities likely predisposed to stress, renders these 

forests highly susceptible to destabilization by native 

insects and pathogens (fig. 9). In historic WWPdominated forests, insects and pathogens probably 

served as a stabilizing agent, removing maladapted 

late seral and climax species relatively early in stand 

development. Most likely, this preserved only the 

best of the climax species and generally encouraged 

dominance of WWP (Harvey and others 1995). Such 

a radical change of endemic processes in a major ecosystem is likely to have far reaching ramifications for 

the productivity, stability, and management (or lack 

thereof) of regional forests (Atkins and others 1999; 

Harvey and others 1995; Monnig and Byler 1992).

Management Implications

Since we are continuing to lose ground with WWP 

and other five-needled pines in the Western United 

States, it is evident that current approaches have not 

been, and will not be, sufficient to reverse losses. 

Discussions from a series of workshops attended by 

regional plant pathologists and geneticists concluded 

that we must at least accomplish the following activities for WWP, and the ecosystems it dominated, to be 
USDA Forest Service RMRS-GTR-208. 2008. 9

restored to even a small portion of its historical range 

(Fins and others 2001):

1) Create more opportunities (and improved rustresistant materials) for artificial regeneration.

2) Create opportunities for “natural” selection, in 

other words, maintain as much as possible of the 

surviving WWP in current forests and provide 

openings for its regeneration in subsequent 

second-generation stands.

3) Develop a broader understanding of alternate host 

ecology and potential resistance.

4) Continue to create opportunities for expanding the 

genetic base for WPBR-resistant stock.

5) Develop a better understanding of the current 

WPBR pathogen potential for adapting to 

changing environments or developing new races.

6) Emphasize coordinated, aggressive management 

with strong Integrated Pest Management 

components (Hagle and others 1989).

7) In non-forested areas, more aggressive planting 

and the use of WWP would at least help maintain 

an extended genetic base for possible future use.

Doing nothing assures that recovery of the western 

white pine ecosystem will not occur for centuries, if 

ever.

Perspectives About the Future 

of Western White Pine

As previously outlined, long-term survival of WWP 

throughout its historical range, except as an occasional 

component of that landscape, may be in jeopardy.

Short-term survival of WWP is not likely in jeopardy (Graham and Jain 1999); however, a substantial 

portion of its genetic breadth may be.

Loss of WWP as a significant component of the 

landscape throughout a substantial portion of its 

historic range has already occurred (Atkins 1999; 

Harvey and others 1995; Monnig and Byler 1992; 

Neuenschwander and others 1999).

Loss of WWP abundantly clear. It has occurred as 

a result of (1) salvage logging of infected trees, (2) 

large trees (infected in the 1940s and 50s) gradually 

succumbing to a combination of the effects of WPBR 

and pine beetles during the 1980s, and (3) lack of opportunities for regeneration.

The largest tree on “white pine drive” in northern 

Idaho died and was removed as a hazard to the public 

in 1999. In 1975, that area was still deserving of the 

name. Subsequently, today, there is hardly a white 

pine to be seen.

Without aggressive intervention sufficient to 

change current trends, the outlook for this magnificent 

tree and, perhaps more importantly, the ecosystem it 

once supported, is obviously dismal. Although current 

efforts have had much success, they have not been 

widespread enough to turn the tide in this battle.

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The Rocky Mountain Research Station develops scientific 

information and technology to improve management, protection, and use of the forests and rangelands. Research is 

designed to meet the needs of the National Forest managers, Federal and State agencies, public and private organizations, academic institutions, industry, and individuals.

Studies accelerate solutions to problems involving ecosystems, range, forests, water, recreation, fire, resource inventory, land reclamation, community sustainability, forest 

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