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Sugar Pine

Pinus lambertiana Douglas

Associations

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Foodplant / parasite
subcortical pycnium of Cronartium ribicola parasitises stem of Pinus lambertiana
Remarks: season: 3-6

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Comments

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The largest species of the genus, Pinus lambertiana also has the longest seed cone in the genus. It is an important timber tree with harvest far exceeding regrowth. It is easily distinguished from P . monticola and P . strobus by its larger cones and thicker cone scales with larger seeds; it is somewhat less reliably distinguished by its leaves, which are slightly wider and more tapering-tipped and have some stomatal lines evident on the abaxial surfaces (the lines not evident in P . monticola and P . strobus ). A "sugary" resin high in cyclitols exudes from the sweet-scented fresh-cut wood.
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
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Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Description

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Trees to 75m; trunk to 3.3m diam., massive, straight; crown narrowly conic, becoming rounded. Bark cinnamon- to gray-brown, deeply furrowed, plates long, scaly. Branches spreading, distal branches ascending; twigs gray-green to red-tan, aging gray, mostly puberulent. Buds cylindro-ovoid, red-brown, to 0.8cm, resinous. Leaves 5 per fascicle, spreading to ascending, persisting 2--4 years, 5--10cm ´ (0.9--)1--1.5(--2)mm, straight, slightly twisted, pliant, blue-green, abaxial surface with only a few lines evident, adaxial surfaces with evident white stomatal lines, margins finely serrulate, apex acuminate; sheath (1--)1.5--2cm, shed early. Pollen cones ellipsoid-cylindric, to 15mm, yellow. Seed cones maturing in 2 years, shedding seeds and falling soon thereafter, often clustered, pendent, symmetric, cylindric before opening, lance-cylindric to ellipsoid-cylindric when open, 25--50cm, yellow-brown, stalks 6--15cm; apophyses somewhat thickened; umbo terminal, depressed, resinous, slightly excurved. Seeds obovoid, oblique apically; body 1--2cm, deep brown; wing broad, 2--3cm. 2 n =24.
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cc-by-nc-sa-3.0
copyright
Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
source
Flora of North America @ eFloras.org
editor
Flora of North America Editorial Committee
project
eFloras.org
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eFloras

Habitat & Distribution

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Montane dry to moist forests; 330--3200m; Calif., Nev., Oreg.; Mexico in n Baja California.
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
source
Flora of North America @ eFloras.org
editor
Flora of North America Editorial Committee
project
eFloras.org
original
visit source
partner site
eFloras

Broad-scale Impacts of Fire

provided by Fire Effects Information System Plants
More info for the terms: basal area, duff, fire severity, fire suppression, forest, fuel, fuel moisture, litter, prescribed fire, severity, tree, wildfire

The Research Project Summary Plant response to prescribed burning with varying season,
weather, and fuel moisture in mixed-conifer forests of California
provides information
on prescribed fire and postfire response
of many plant community species including sugar pine.

Near the Plumas National Forest, prescribed fire in a mixed-conifer-California
black oak forest with a sugar pine component successfully reduced fuel load.
When a wildfire burned through the site previously burned under prescription,
fire severity and fire suppression costs were less compared to adjacent land
where fire had been excluded [27]. For further information on this study, see the Research Paper by Moghaddas [27].

A fall prescribed fire in the Tharp Creek Watershed of Sequoia National Park
produced 17.2% and 11.7% average annual sugar pine mortality on 2 white fir-mixed
conifer sites monitored for 5 years after fire. Mortality was concentrated  in the
subcanopy. The fire burned from 23 to 26 October 1990. Relative humidity during
the day was 21% to 30% and at night was 30% to 40%. Fuel moisture levels in the
litter and duff averaged 28%. For 100-hour and 1,000-hour fuels, moisture levels
were 14% and 64%, respectively. At the time of ignition, air temperatures were
50 to 61 °F (10-16 °C) and winds were calm. The fire was a combination of backing and strip headfires with flame lengths of 0.16 to 7.9 feet (0.05-2.4 m).
One-hour, 10-hour, and 100-hour fuels were reduced by 96%, 77%, and 60%, respectively.
Tree (≥4.6 feet (1.4 m)) mortality was evaluated before and after fire as
well as from an unburned reference site. Basal area (m²/ha) changes were also
monitored before and after the fire. Mean annual percent change in sugar pine basal
area increased by an average of 0.17% and 1.39% on the 2 burned sites before the fire
compared to the control site.  From 1989 to 1994 (includes 1 year of prefire data),
sugar pine basal area was reduced 4.28% to 15.67% on the burned sites compared to
the control [28]. For more information, see the entire Research Paper by Mutch and
Parsons [28].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Common Names

provided by Fire Effects Information System Plants
sugar pine
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Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Cover Value

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More info for the term: cover

Sugar pine is used for cover by wildlife.  Early in sugar pine
development, large mammals use dense stands as hiding and thermal cover.
Mature trees are used by arboreal species such as birds, squirrels, and
other small mammals.  Old-growth sugar pine is prime habitat for cavity
nesters such as woodpeckers and owls [16].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Description

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Sugar pines may live 400 to 500 years and are second only to giant
sequoia (Sequoia gigantea) in total volume.  A record sugar pine in
California measured 216 feet (66 m) tall and 122 inches (310 cm) in
d.b.h.  Trees up to 250 feet (76 m) tall and 10 feet (3 m) in diameter
have been reported.  Mature sugar pine cones are among the largest of
all conifers, averaging 12 inches (30 cm) in length, and can reach 22
inches (56 cm) long.  Its needles are 3 inches (7.5 cm) long and have
five to a cluster.  Sugar pines pyramidal crown has whorls of horizontal
branches with several conspicuously longer than others.  Its sap
contains a sugary substance [7,16,21].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Distribution

provided by Fire Effects Information System Plants
Sugar pine extends from the western slope of the Cascade Range in
north-central Oregon to the Sierra San Pedro Mártir in Baja California.
Its distribution is almost continuous through the Klamath and Siskiyou
mountains and on western slopes of the Cascade Range and Sierra Nevada.
Smaller and more disjunct populations are found in the Coast Range of
southern Oregon and California, Transverse and Peninsula ranges of
southern California, and east of the Cascade and Sierra Nevada crests.
Its southern extremity is an isolated population high on a plateau in
the Sierra San Pedro Mártir in Baja California, Mexico.  Over 80 percent
of its distribution is in California [16,21].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Fire Ecology

provided by Fire Effects Information System Plants
More info for the term: fire regime

Sugar pine is very resistant to low- to moderate-severity fires.  It has
adapted a thick, fire-resistant bark and open canopy that retards aerial
fire spread.  Young sugar pine seedlings prefer bare mineral seedbeds
[2,3].

FIRE REGIMES :
Find fire regime information for the plant communities in which this
species may occur by entering the species name in the FEIS home page under
"Find FIRE REGIMES".
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Fire Management Considerations

provided by Fire Effects Information System Plants
Prescribed burning has been found to be an effective management
treatment that will destroy infected stands of sugar pine where dwarf
mistletoe and other diseases have rendered stands unmerchantable [1].
Dead sugar pine is susceptible to blue stain fungus in the sapwood;
however, the heartwood is very durable.  Salvagable trees may be found
up to 17 years after being killed by fire [15].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Growth Form (according to Raunkiær Life-form classification)

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More info on this topic.

More info for the term: phanerophyte

Phanerophyte
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat characteristics

provided by Fire Effects Information System Plants
More info for the term: mesic

Sugar pine is found on a variety of sites from moist, steep, north- and
east-facing slopes, to more mesic, south-facing slopes.  The fuels under
sugar pine are generally heavy with deep soils.

Climate:  Temperature and precipitation vary widely throughout the range
of sugar pine.  The general weather pattern consists of hot, dry
summers and cool, wet winters.  Precipitation during July and August is
usually less than 1 inch (2.5 cm) per month and summertime relative
humidities are low.  Most precipitation occurs between November and
April, mostly in the form of snow at middle elevations.  Total
precipitation varies from 33 to 69 inches (83-173 cm) per year [16].

Soils and topography:  Soil parent material include rocks of volcanic,
granitic, and sedimentary origin.  Soils formed from peridotite or
serpentinite typically support sugar pine stands of inferior growth and
quality.  The most extensive soils supporting sugar pine are
well-drained, moderately to rapidly permeable, and slightly acidic to
neutral pH (7.0).  Best development of sugar pine is on mesic soils with
sandy to clayey loam textures.  Much of the terrain occupied by sugar
pine is steep and rugged.  Sugar pines are equally distributed on all
aspects at lower elevations but grow best on warm exposures as elevation
increases.  Optimum growth occurs on gentle terrain at middle elevations
[16].

Elevation:  Sugar pine ranges from near sea level in the Coast Range to
more than 10,000 feet (3,000 m) in the Transverse Range.  Elevational
limits increase with decreasing latitude.  Typical elevational ranges
are as follows [16]:
                   Cascade Range:  1,100 to  5,400 feet (335-1,645 m)  
                   Sierra Nevada:  2,000 to  7,500 feet (610-2,285 m)
         Sierra San Pedro Mártir:  7,056 to  9,100 feet (2,150-2,775 m)
Transverse and Peninsular Ranges:  4,000 to 10,000 feet (1,220-3,000 m)
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Cover Types

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This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

   207  Red fir
   211  White fir
   229  Pacific Douglas-fir
   231  Port-Orford-cedar
   232  Redwood
   234  Douglas-fir - tanoak - Pacific madrone
   244  Pacific ponderosa pine - Douglas-fir
   246  California black oak
   247  Jeffrey pine
   249  Canyon live oak
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Ecosystem

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This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

   FRES20  Douglas-fir
   FRES21  Ponderosa pine
   FRES26  Lodgepole pine
   FRES27  Redwood
   FRES28  Western hardwoods
   FRES34  Chaparral - mountain shrub
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Plant Associations

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More info on this topic.

This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the terms: forest, shrub

   K002  Cedar - hemlock - Douglas-fir forest
   K005  Mixed conifer forest
   K006  Redwood forest
   K007  Red fir forest
   K008  Lodgepole pine - subalpine forest
   K010  Ponderosa shrub forest
   K034  Montane chaparral
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Immediate Effect of Fire

provided by Fire Effects Information System Plants
Sugar pine is rated as intermediate in fire tolerance.  Young sugar
pines are susceptible to low- to high-severity fires.  Mature trees can
survive most fires, suffering only bole scorch.  Sugar pine
susceptibility to secondary attack by insects and disease following fire
is rated as low [3].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Importance to Livestock and Wildlife

provided by Fire Effects Information System Plants
Birds and mammals use sugar pine as a source of food and shelter.
Douglas' squirrels and white-headed woodpeckers have been noted to
occupy sugar pine trees [16].
license
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Key Plant Community Associations

provided by Fire Effects Information System Plants
More info for the terms: codominant, forest, mesic, woodland

Sugar pine usually occurs in mixed-conifer forest stands with a wide
variety of overstory associates including ponderosa and Jeffrey pine
(Pinus ponderosa and P. jeffreyi), California red fir (Abies magnifica),
white fir (A. concolor), noble fir (A. procera), and Douglas-fir
(Pseudotsuga menziesii) [4,21].  In southern California, sugar pine is
characteristically found in vegetation types of the woodland and
timberland chaparral zones.  Canyon live oak (Quercus chrysolepis) is
found with sugar pine on more mesic sites, while at higher elevations
sugar pine occurs with mountain whitethorn (Ceanothus cordulatus), Parry
manzanita (Arctostaphylos parryana var. pinctorum), and bush chinquapin
(Chrysolepsis sempervirens) [14].

Publications listing sugar pine as a codominant species in plant
vegetation types (vts) or community types (cts) are listed as follows:

Area                   Classification                       Authority
----                   --------------                       ---------
s CA                    forest (vts)                        Horton 1960
s CA                    forest (cts)                        Thorne 1977
  CA                    forest (cts)                        Thorne 1976
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Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Life Form

provided by Fire Effects Information System Plants
More info for the term: tree

Tree
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Management considerations

provided by Fire Effects Information System Plants
More info for the term: cone

Sugar pine is planted on a vast scale in Oregon and California, and also
has been tried in several countries around the world.  Large-scale
plantings, however, are few due to establishment difficulties and
restrictive site requirements for good growth [21].  Sugar pine does not
self-prune; therefore, high-quality clear-lumber requires the pruning of
lower limbs.  It is the most tolerant to oxidant air pollution among its
coniferous associates [8,16].

Disease:  Sugar pine is highly susceptible to white pine blister rust
caused by the fungus Cronartium ribicola.  Among commercially important
North American white pines, sugar pine is the most susceptible to this
disease.  Infected seedlings and young trees are inevitably killed by
cankers girdling the main stem.  Incidence and intensity of infection on
sugar pine are highest in Oregon and northern California and become
progressively less to the south, as the climate becomes warmer and
drier.  Dwarf mistletoe (Arceuthobium californicum) may seriously damage
infected trees, but spread is slow and can be controlled by sanitation
cutting [13,16,21].

Insects:  The most damaging insect threatening sugar pine is the
mountain pine beetle (Dendroctonus ponderosae).  During periods of
drought, other insects such as the red turpentine beetle (D. valens) and
California flathead borer (Melanophila californica) usually attack
unhealthy trees and those under moisture stress.  The sugar pine cone
beetle (Conophthorus lambertianae) is extremely destructive to
developing second-year cones [5,16].

Animals:  Small mammals such as pocket mice, jumping mice, chipmunks,
and ground squirrels forage on young seedlings, thus reducing
regeneration on disturbed sites [3].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Occurrence in North America

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     CA  NV  OR  MEXICO
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Other uses and values

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Native Americans used the pitch from sugar pine to repair canoes and to
fasten arrowheads and feathers to shafts [2].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Palatability

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Sugar pine is considered low in palatability to livestock and wildlife.
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Phenology

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Seasonal growth durations of sugar pine at various elevations in the Sierra
Nevada are as follows [11]:

                   Height           Radial
                   Growth*          Growth
                   ------           ------
Start (days)**       146              107
Start (date)       May 26         April 17
Length (days)         51              129
Rapidity (days)       15               46

* An 8-year average.
** Number of days from January 1.
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Post-fire Regeneration

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More info for the term: seed

   off-site colonizer; seed carried by wind; postfire years 1 and 2
   off-site colonizer; seed carried by animals or water; postfire yr 1&2
license
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Regeneration Processes

provided by Fire Effects Information System Plants
More info for the terms: basal area, cone, epigeal, fresh, litter, monoecious, seed, stratification

Sugar pine does not sprout, but young trees can be rooted from cuttings.
Its primary regeneration strategy is via seed [3,16].

Flowering and fruiting:  Sugar pine is monoecious.  Reproductive buds
are set in July and August, but are not discernible until late the next
spring.  Time of pollination ranges from late May to early August,
depending on elevation.  Female strobili are approximately 1 to 2 inches
(2.5-5.0 cm) long when pollinated and may double in size by the end of
the growing season.  Fertilization occurs the following spring,
approximately 12 months after pollination.  Dates of cone opening range
from mid-August at low elevations to early October at high elevations.
Sugar pine does not become a good cone producer until it has attained a
diameter of about 30 inches (75 cm) or is about 150 years old [2,16].

Seed production and dissemination:  Mature trees produce large amounts
of seeds, averaging up to 150 seeds per cone.  In good crop years, the
proportion of sound seeds is usually high (67 to 99 percent) but in
light crop years can fall as low as 28 percent.  Seed shed may begin in
late August at low elevations and at higher elevations is usually
complete by the end of October.  Seeds are large and heavy, averaging
2,100 seeds per pound (4,630/kg).  Seeds are not dispersed great
distances by wind, and 80 percent fall within 100 feet (30 m) of the
source.  Birds and small mammals aid in seed dissemination [16].

Seedling development:  Sugar pine seeds may lie dormant, but dormancy
can be broken by a 60 to 90 day stratification.  Fresh seed may
germinate with a 90 percent success rate if adequately ripened, cleaned,
and stratified.  Losses due to unprepared seedbeds, drought, insects,
and rodents may be high.  Germination is epigeal.  Seedlings rapidly
grow a deep taproot when seeds germinate on mineral soil.  Seedlings
will germinate on both litter and bare mineral soil, but development is
slow under shade conditions.  After 2 years, taproots range from 22 to
40 inches (56-102 cm) deep.  Planting sugar pine has met with some
failure.  A low drought tolerance may be the determining factor.  Sowing
stratified seed in February or March extends the growing season and
produces healthy seedlings of plantable size in one season [4,16].

Growth and yield:  Early growth of sugar pine is slow compared to
ponderosa pine but increases rapidly in the pole stage and continues
through maturity.  On favorable sites, growth increments in basal area
of 2.5 percent or more can be sustained for up to 100 to 150 years.  The
best growth can be found between 4,500 to 6,000 feet (1,370-1,830 m) in
the central Sierra Nevada, between the American and San Joaquin Rivers.
Sugar pine is semitolerant to shade and may exhibit poor growth if
seedlings are enclosed by brush.  Sugar pine is a deep-rooted species
that is not susceptible to windthrow [9,16,21].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Regional Distribution in the Western United States

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This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

   1  Northern Pacific Border
   3  Southern Pacific Border
   4  Sierra Mountains
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Successional Status

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More info for the terms: climax, forest, natural

Sugar pine is primarily an early-seral to seral species.  It is
rarely found in pure stands.  When sugar pine is found to be the dominant
species in old-growth stands, it most often was dominant to begin with
or released by natural causes.  White fir would usually be the climax
species in mixed conifer forest in the absence of any natural
disturbances.  When disturbance does occur, it creates gaps in which
sugar pine is well adapted to grow [3,4,16,25].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Taxonomy

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The currently accepted scientific name of sugar pine is Pinus
lambertiana Dougl. [24]. There are no recognized subspecies, varieties,
or forms.
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Wood Products Value

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High-grade sugar pine lumber is sought after for its dimensional
stability and workability.  The wood is light and resists deformity.  It
is easily milled and is favored for molding, window and door frames,
window sashes, doors, and other special products like piano keys and
organ pipes [16].
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bibliographic citation
Habeck, R. J. 1992. Pinus lambertiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Associated Forest Cover

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Sugar pine is a major timber species at middle elevations in the Klamath and Siskiyou Mountains, Cascade, Sierra Nevada, Transverse, and Peninsula Ranges. Rarely forming pure stands, it grows singly or in small groups of trees. It is the main component in the forest cover type Sierra Nevada Mixed Conifer (Society of American Foresters Type 243) (10) generally comprising 5 to 25 percent of the stocking. It is a minor component in 10 other types:

207 Red Fir
211 White Fir
229 Pacific Douglas-Fir
231 Port-Orford-Cedar
232 Redwood
234 Douglas-Fir-Tanoak-Pacific Madrone
244 Pacific Ponderosa Pine-Douglas-Fir
246 California Black Oak
247 Jeffrey Pine
249 Canyon Live Oak

In the northern part of its range, sugar pine is commonly associated with Douglas-fir (Pseudotsuga menziesii), ponderosa pine (Pinus ponderosa), grand fir (Abies grandis), incense-cedar (Calocedrus decurrens), western hemlock (Tsuga heterophylla), western redcedar (Thuja plicata), Port-Orford-cedar (Chamaecyparis lawsoniana), tanoak (Lithocarpus densiflorus), and Pacific madrone (Arbutus menziesii). In the central part it is associated with ponderosa pine, Jeffrey pine (Pin us jeffreyi), white fir (Abies concolor), incense-cedar, California red fir (A. magnifica), giant sequoia (Sequoiadendron giganteum), and California black oak (Quercus kelloggii). Farther south, the usual associates are Jeffrey pine, ponderosa pine, Coulter pine (Pinus coulteri), incense-cedar, white fir, and bigcone Douglas-fir (Pseudotsuga macrocarpa). At upper elevations Jeffrey pine, western white pine (Pinus monticola), California red fir, and lodgepole pine (P. contorta) grow with sugar pine. In the Sierra San Pedro Martir, Jeffrey pine and white fir are the main associates.

Common brush species beneath sugar pine include greenleaf manzanita (Arctostaphylos patula), deerbrush (Ceanothus integerrimus), snowbrush (C. velutinus), mountain whitethorn (C. cordulatus), squawcarpet (C. prostratus), bearclover (Chamaebatia foliolosa), bush chinkapin (Castanopsis sempervirens), bitter cherry (Prunus emarginata), salal (Gaultheria shallon), coast rhododendron (Rhododendron californicum), and gooseberries and currants in the genus Ribes (11). From a silvicultural standpoint, Ribes spp. are especially important because they are alternate hosts to the white pine blister rust fungus (Cronartium ribicola). At least 19 different species grow in the Mixed Conifer Type, of which the Sierra gooseberry (Ribes roezlii) is most prevalent on more xeric, upland sites, and the Sierra currant (R. nevadense) on more mesic sites (35).

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Climate

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Temperature and precipitation vary widely throughout the range of sugar pine. For equivalent latitudes, temperature decreases and precipitation increases with elevation, and for equivalent elevations, temperature increases and precipitation decreases from north to south. Patterns unifying this variability are relatively warm, dry summers and cool, wet winters. Precipitation during July and August is usually less than 25 mm (1 in) per month, and summertime relative humidities are low. Although water stored in snowpacks and soils delays the onset and shortens the duration of summer drought, evaporative stress often becomes great enough to arrest growth in the middle of the season (15). Most precipitation occurs between November and April, as much as two-thirds of it in the form of snow at middle and upper elevations (26). Within its natural range, precipitation varies from about 840 to 1750 mm (33 to 69 in). Because winter temperatures are relatively mild and seldom below freezing during the day, considerable photosynthesis and assimilation are possible during the dormant season, at least partially offsetting the effects of summer drought (15).

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Damaging Agents

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The pathology of sugar pine is dominated by white pine blister rust, caused by Cronartium ribicola, a disease serious enough to severely limit natural regeneration in areas of high hazard, and thereby alter successional trends. Among commercially important North American white pines, sugar pine is the most susceptible. Infected seedlings and young trees are inevitably killed by cankers girdling the main stem.

Blister rust was introduced into western North America shortly after the turn of the century at a single point on Vancouver Island and has since spread eastward throughout the Inland Empire and south through the Cascade, Klamath, North Coast, and Sierra Nevada Ranges. It has not yet been found in the Transverse or Peninsular Ranges of southern California, even though alternate host species are abundant there. Within the range of sugar pine, conditions for infection are not nearly so uniform as for western white pine in the Inland Empire. Incidence and intensity of infection on sugar pine are highest in Oregon and northern California and become progressively less to the south, as climate becomes warmer and drier. Within any area, however, hazard varies widely and depends on local site conditions. These are complex, but two of the most important factors are the duration of moisture retention on foliage following rain, fog, or dew, and the distribution and density of the alternate hosts, currant and gooseberry bushes (Ribes spp.). Thus, cool north slopes are more hazardous than warm south slopes, and relatively humid stream bottoms and lakesides are more hazardous than upland sites. In the Cascade Range and Sierra Nevada of northern California, infection averaged two to three times higher near stream bottoms than on adjacent slopes (4).

Attempts to control blister rust by chemical therapy or eradicating alternate hosts have been abandoned as impractical and ineffective. Except on highly hazardous sites, sugar pine in natural stands can be effectively managed by judiciously selecting leave trees with cankers relatively far from the bole and by pruning cankers in the lower crown (4).

Plantations are a much more serious problem. The microenvironmental changes on a site following clearcutting-including dew formation on foliage and the rapid regeneration of alternate host Ribes spp. greatly augment the probability of rust intensification and spread on both hosts. Uniform age and stocking make sugar pine plantations vulnerable to nearly total destruction for 20 years or longer. Genetically resistant sugar pines in mixture with other conifers offer the most promising solution.

Dwarf mistletoe (Arceuthobium californicum) may seriously damage infected trees by reducing growth in height, diameter, and crown size, and predisposing weakened trees to attack by bark beetles. Extending throughout the range of sugar pine, except for isolated stands in Nevada, the south Coast Ranges of California, and Baja California, this mistletoe was found in only 22 percent of the stands examined and on only 10 percent of the trees in those stands. Spread is slow and can be controlled by sanitation cutting (20,42).

A needle cast caused by Lophodermella arcuata is occasionally and locally damaging. Root diseases caused by Armillaria mellea, Heterobasidion annosum, and Verticicladiella wageneri are capable of killing trees of all ages and sizes but, though widespread, are usually at endemic levels. Several trunk and butt rots attack sugar pine but are usually confined to mature and overmature trees (2,21).

Several root and damping-off pathogens cause severe damage to sugar pine in nurseries, with annual losses up to 50 percent (45). In approximate order of importance, these are Fusarium oxysporum, Macrophomina phaseoli, and species of Pythium, Phytophthora, and Rhizoctonia. In addition to causing direct losses in the nursery, these diseases may reduce field survival of planted seedlings in more stressful environments by causing stunting and chlorosis. Nursery fumigation controls most of the organisms involved but is least effective on Fusarium. A simple and promising alternative control method is early sowing of stratified seed. Soil temperatures in late winter and early spring permit seed germination and root development but are still cool enough to inhibit fungal growth.

Sugar pine hosts many different insects, but the mountain pine beetle (Dendroctonus ponderosae) is of overwhelming importance. This insect can cause widespread mortality, often killing large groups of trees (48). Several other bark-feeding insects contribute directly or indirectly to mortality in sugar pines, particularly after periods of drought. Death results from predisposing trees to mountain pine beetle. The red turpentine beetle (Dendroctonus valens) is usually restricted to small areas near the root crown but during drought may extend two or more meters up the bole, destroying the entire cambium. The California flatheaded borer (Melanophila californica) usually attacks decadent and unhealthy trees, but trees under heavy moisture stress are also vulnerable. The California fivespined ips (Ips paraconfusus) is only capable of penetrating thin bark in sugar pine. Small trees are often killed, but large trees only top-killed (16).

The sugar pine cone beetle (Conophthorus lambertianae) can be extremely destructive to developing second-year cones, destroying up to 75 percent of the crop in some years. Since stunted cones are apparent by mid-June, the extent of the crop loss can be assessed well before cone collection. The sugar pine scale (Matsucoccus paucicicatrices) occasionally kills foliage and branches, predisposing trees to bark beetle attack. The dead "flags" resulting from heavy attack mimic advanced symptoms of white pine blister rust. Occasionally, the black pineleaf scale (Nuculaspis californica) defoliates sugar pine at midcrown, weakening the tree. These scale attacks are often associated with industrial air pollution or heavy dust deposits on foliage (16).

Among its coniferous associates, sugar pine is the most tolerant to oxidant air pollution (34), while intermediate in fire tolerance (39) and frost tolerance (43,44). It is less tolerant of drought than most companion species with which it has been critically compared, including knobcone (Pinus attenuata) and Coulter pines (50,51), ponderosa pine, Douglas-fir, incense-cedar, and grand fir (40).

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Flowering and Fruiting

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Sugar pine is monoecious. Reproductive buds are set in July and August but are not discernible until late in the next spring. Time of pollination ranges from late May to early August, depending on elevation, and to a lesser extent on latitude.

Female strobili are 2.5 to 5.0 cm (1 to 2 in) long at time of pollination and double in size by the end of the growing season. Fertilization of eggs by male gametes takes place late the following spring, about 12 months after pollination. By this time, the seed is at its final size with a fully developed coat. Conelet elongation continues during the second season until maturation in late summer. Mature sugar pine cones are among the largest of all conifers, averaging 30 cm (12 in) and ranging up to 56 cm (22 in) long. Dates of cone opening range from mid-August at low elevations to early October at high elevations (12,19,32).

Cone production starts later and is less prolific in sugar pine than in its associates. During a 16-year study in the central Sierra, fewer than 5 percent of sugar pines less than 20 cm (8 in) in d.b.h., and 50 percent less than 31 cm (12 in) in d.b.h., produced cones. Of trees 51 cm (20 in) or more, 80 percent produced cones, and dominant trees produced 98 percent of the total. Intervals between heavy cone crops averaged 4 years and ranged from 2 to 7 (12).

Loss of sugar pine cones is heavy; the probability of a pollinated conelet developing to maturity is only 40 to 50 percent. Predation by the sugar pine cone beetle (Conophthorus lambertianae) can cause up to 93 percent loss. Douglas squirrels and white-headed woodpeckers also take a heavy toll (7,11,17).

Spontaneous abortion of first-year conelets is high. Observations of control-pollinated trees in the Klamath Mountains showed that 19 percent of female strobili were lost 5 to 12 weeks after bagging, with no obvious signs of insect or pathogen-caused damage (41). The amount of abortion varied from 15 to 85 percent among trees, for both bagged and unbagged strobili. Since this pattern was consistent in successive years, a genetic cause was suggested.

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Genetics

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Sugar pine is one of the more genetically variable members of the genus. Average heterozygosity of specific genes coding for seed proteins (isozymes) was 26 percent, a value near the upper range (0 to 36 percent) of pines studied so far (6). How adaptive variation is distributed over the range of environments encountered in over 14° of latitude and 2000 m (6,560 ft) of elevation is largely unknown, however, because of a lack of field data from provenance or progeny tests.

In a 3-year nursery trial, pronounced differences in height and diameter growth were found among seedlings of five seed sources sampled along an elevational transect on the west slope of the Sierra Nevada (18). The fastest growing seedlings were from the lower-middle elevation (1100 m or 3,595 ft) and were twice the height of those from the highest elevation (2195 m or 7,200 ft). Except for the source from the lowest elevation (770 m or 2,525 ft), which ranked second, growth varied inversely with elevation. Elevation of the seed source accounted for 52 percent of the total variance among seedlings, and the component of variance for families within stands was a substantial 16 percent. More comprehensive nursery trials, of families from seed parents ranging from southern California to southern Oregon, showed similar trends (27). Greatest growth was expressed in seedlings from intermediate elevations in the central Sierra Nevada, a result consistent with observations in natural stands. Thus, genetic adaptation to climatic variables associated with elevation is clearly evident in sugar pine, requiring a close match between seed source and planting site in artificial regeneration. The degree of variability expressed among progenies of different seed parents within seed collection zones indicates that selection for rapid early growth should be effective.

Resistance to white pine blister rust is strongly inherited, and three different kinds have been recognized (29). A rapid, hypersensitive reaction to invading mycelium is conditioned by a dominant gene. This gene, which occurs at variable but relatively low frequencies throughout the range of sugar pine, is highly effective against most sources of inoculum. A race of blister rust capable of overcoming this gene was discovered in a plantation in the Klamath Mountains (30), but evidently had not spread from this site 10 years after it was found (31). In certain families, another kind of resistance is expressed by slower rates of infection and mortality, fewer infections per tree, and by a higher rate of abortion of incipient infections. This "slow rusting" is apparently inherited quantitatively and, while less dramatic than single gene resistance, may be more stable to variation in the pathogen in the long term. Probably two or more generations of selection and breeding will be necessary to accumulate enough genes in parental stock to make this kind of resistance usable in commercial silviculture. A third kind of resistance is age-dependent. In common garden tests, infection among grafted clones from mature trees ranged from 0 to 100 percent, yet offspring from the apparently resistant clones were fully susceptible. Although not understood, the mechanisms and inheritance of mature tree resistance are very strong and could play a significant role in stabilizing resistance over a rotation. Since all three kinds of resistance are inherited independently, there is a real promise for an enduring and well-buffered genetic control of this most destructive disease.

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Growth and Yield

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Veteran sugar pines often reach great size. Large trees have commonly scaled 114 to 142 m³ (20,000 to 25,000 fbm, Scribner log rule), with a record of 232 m³ (40,710 fbm). A "champion," located on the North Fork of the Stanislaus River in California, measured 65.8 m (216 ft) tall and 310 cm (122 in) in d.b.h., but trees up to 76 m (250 ft) tall have been reported (11,36). These and previous champions of this century are dwarfed by the first sugar pine measured by David Douglas and described in his diary (37): "Three feet from the ground, 57 feet 9 inches in circumference; 134 feet from the ground, 17 feet 5 inches; extreme length 215 feet."

Early growth of sugar pine is slow compared with ponderosa pine, but growth rates accelerate in the pole stage and are sustained for longer periods than those of common associates. Consequently, sugar pines are usually the largest trees, except for giant sequoia, in mature and old-growth stands. On better sites annual growth increments in basal area of 2.5 percent and more can be sustained up to stem diameters of 76 to 127 cm (30 to 50 in) or for 100 to 150 years (11). Growth of sugar pine is best between 1370 and 1830 m (4,500 and 6,000 ft) in the central Sierra Nevada, between the American and San Joaquin Rivers.

In young mixed conifer stands, sugar pine often constitutes a relatively small proportion of the total basal area but contributes disproportionately to growth increment. On the El Dorado National Forest in the western Sierra Nevada, in stands ranging in age from 50 to 247 years, the sugar pine component was only 6 to 7 percent (range: 3 to 14 percent) of the average basal area, but its average annual basal area growth was 11.3 percent (range: 2 to 35 percent) of the stand total. A similar relationship was found on the Plumas National Forest in the northern Sierra Nevada: in stands from 58 to 95 years old, average basal area of sugar pine was 7 percent (3 to 16), but 10-year growth was more than 12 percent (6 to 19). Ten-year volume increment in mixed conifer stands from 40 to 80 years old was greater for sugar pine than for Douglas-fir, white fir, ponderosa pine, and incense-cedar in each of five basal area categories (9). Mean increment for sugar pine was 4.1 percent, compared to 3.1 percent for all others.

Yields of sugar pine are difficult to predict, because it grows in mixes of varying proportion with other species. In the old-growth forest, the board foot volume of sugar pine was 40 percent of total in stands dominated by ponderosa pine and sugar pine. In exceptional cases on very small areas, yields were 2688 m³/ha (192,000 fbm/acre) (11). Yield tables for young growth are based on averages for all commercial conifers and assume full stocking (8). The data base is limited, so the tables are at best a rough guide. Realistically, yields may reach 644 m³/ha (46,000 fbm/acre) in 120 years on medium sites, and Up to 1190 m³/ha (85,000 fbm/acre) in 100 years on the best sites, with intensive management (11).

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Reaction to Competition

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Sugar pine tolerates shade better than ponderosa pine but is slightly less tolerant than incense-cedar and Douglas-fir and much less so than white fir (14). A seral species, it becomes less tolerant with age, and overtopped trees decline unless released (11). Thus, dominant sugar pines in old-growth stands were probably dominant from the start, or released by natural causes early in life. White fir would usually be the climax species in mixed conifer forests in the absence of any natural disturbance; however, fire, insects, disease, and other agents are natural and pervasive features of these forests. Such disturbances frequently cause gaps, in which the relatively tolerant sugar pine is adapted to grow (14). For these reasons, sugar pine is often adapted to regenerate in a shelterwood silvicultural system (33).

Competition from brush severely retards seedling establishment and growth. Only 18 percent of seedlings starting under brush survived over a period of 18 to 24 years, and after 10 years the tallest seedlings measured were only 29 cm (11.4 in). Given an even start with brush, however, seedlings can compete successfully (11).

Light shelterwoods can protect seedlings of sugar pine and white fir against frost, which seldom affects ponderosa and Jeffrey pines, and provide them with a competitive advantage because of their greater tolerance to shade (13,43,44). On the other hand, young sugar pines stagnate beneath an overstory and in competition with root systems of established trees or brush. But because they respond well to release, the basal area increment of sugar pines is often double that of companion species after heavy thinnings (33). Thus, skill in the amount and timing of overstory removal is a key factor in successful silvicultural management of sugar pine.

Sugar pine does not self-prune early, even in dense stands, and mechanical pruning is necessary to ensure clear lumber of high quality.

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Rooting Habit

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Sugar pine develops a deep taproot at an early age.

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Seed Production and Dissemination

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Mature trees produce large amounts of sound seeds. In a study of 210 trees in 13 stands in the central and northern Sierra Nevada, the average number of sound seeds per cone was 150, with individual trees ranging from 34 to 257. Higher numbers of seeds per cone (209 to 219) have been reported, but whether the count was based on sound or total seeds was not specified. In good crop years, the proportion of sound seeds is usually high (67 to 99 percent) but in light crop years can fall as low as 28 percent (7,12).

Cones are ripe and start to open when their color turns light brown and specific gravity (fresh weight basis) drops to about 0.62. Seed shed may begin in late August at low elevations and at higher elevations is usually complete by the end of October (11).

Seeds are large and heavy, averaging 4,630 seeds per kilogram (2,100/lb). Since their wings are relatively small for their size, seeds are not often dispersed great distances by wind, and 80 percent fall within 30 m (100 ft) of the parent tree. Birds and small mammals may be an important secondary mechanism of dispersal, even though they consume most of the seeds they cache. In good seed years, large amounts of seed fall, with estimates ranging from 86,500 to more than 444,800/ha (35,000 to 180,000/acre) in central Sierra Nevada stands (11,32).

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Seedling Development

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Sugar pine seeds show dormancy, which can be readily broken by stratification for 60 to 90 days or by removal of the seed coat and inner papery membrane surrounding the seed. Germination of fresh seed is uniformly rapid and high, exceeding 90 percent if adequately ripened, cleaned, and stratified. Viability may decline rapidly with time in storage at temperatures above freezing, but deep-frozen seed maintains viability much longer (1,32,47).

On unprepared seed beds, seed-to-seedling ratios are high (244 to 483). Soil scarification reduced the ratio to 70 in one case, and scarification with rodent poisoning dropped it to 38 in another (12).

Seedling losses are continual and only 20 to 25 percent of the initial germinants may survive as long as 10 years. Drought may kill up to half of the first-year seedlings. Cutworms and rodents, which eat seeds still attached to seedling cotyledons, also take their toll (11,12). Seedlings infected by blister rust rarely survive more than a few years.

Germination is epigeal (32). Seedlings rapidly grow a deep taproot when seeds germinate on bare mineral soil. In one comparison, taproots penetrated to an average depth of 43 cm (17 in) on a bare sandy soil, but only half as deep when the soil was overlain with duff (11). Lateral roots develop near and parallel to the soil surface, often growing downward some distance from the stem. In heavier, more shallow soils, laterals are often larger than taproots. During the second season, laterals commonly originate on the lower taproot and occupy a cone of soil which has its base at the tip of the taproot. After 2 years on three different soil types in Oregon, the taproots of natural sugar pine seedlings ranged from 56 to 102 cm (22 to 40 in), were significantly deeper than those of Douglas-fir and grand fir, but shorter than those of ponderosa pine and incense-cedar. Lengths of main lateral roots showed the same species differences. Top-to-root ratios for sugar pine ranged from 0.17 to 0.28 (length) and from 1.33 to 1.60 (dry weight) (46).

Seasonal shoot growth starts later and terminates earlier in sugar pine than in its usual conifer associates, except white fir. At middle elevations in the central Sierra Nevada, shoot elongation begins in late May, about 2 weeks after ponderosa pine and a month before white fir, and lasts about 7 weeks. Radial growth begins about 6 weeks earlier than shoot growth and extends throughout the summer (11).

Planting of sugar pine has not been so easy or successful as for some of the yellow pines. Although reasons for the many recorded failures are often complex, lower drought tolerance may be one of the factors. During natural regeneration, the ability of sugar pine seedlings to avoid summer drought by rapidly growing a deep taproot largely compensates for the relative intolerance of tissues to moisture stress (38).

To survive the first summer after planting, seedlings must have the capacity to regenerate vigorous new root systems. For other western conifers, root growth capacity is conditioned by particular combinations of nursery environment and time in cold storage after lifting; these requirements are species and seed-source specific (22,24,38). Although patterns of root growth capacity have not been worked out for sugar pine, it is clear that amounts of root growth are substantially less for sugar pine than for its associates (23).

Early top growth of sugar pine is not so rapid as that of western yellow pines, and 1-year stock is too small for planting when seed is sown in May, for years the tradition in California nurseries. Root diseases, to which young sugar pines are unusually vulnerable, can compound the problem by weakening seedlings that survive, thus reducing their chances of establishment on the site. Sowing stratified seed in February or March extended the growing season and produced healthy seedlings of plantable size in one season (23). A more expensive alternative to bareroot stock that holds some promise is containerized seedlings grown under accelerated growth regimes (28).

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Soils and Topography

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Sugar pine grows naturally over a wide range of soil conditions typically associated with conifer-hardwood forests. Soil parent materials include rocks of volcanic, granitic, and sedimentary origin and their metamorphic equivalents and are usually not of critical importance. Soils formed on ultrabasic intrusive igneous rocks such as peridotite and serpentinite, however, have low calcium-to-magnesium ratios and usually support open conifer stands of inferior growth and quality. Nevertheless, sugar pine is often the dominant conifer on the more mesic of these sites (39,40).

Because site productivity is a function of several environmental variables-edaphic, climatic, and biotic-it is difficult to relate parent material groups or particular soil series with specific productivity classes, especially when they span wide ranges of elevation and latitude. Other factors being equal, the main edaphic influences on conifer growth are soil depth and texture, permeability, chemical characteristics, and drainage and runoff properties (5).

The most extensive soils supporting sugar pine are well drained, moderately to rapidly permeable, and acid in reaction. Soils derived from ultrabasic rocks are very slightly acid to neutral (pH 7.0). In general, acidity increases with soil depth. Several edaphic properties are influenced by the degree of soil profile development. Soil porosity, permeability, and infiltration rate decrease with more developed profiles, while water-holding capacity, rate of run-off, and vulnerability to compaction increase.

Sugar pine reaches its best development and highest density on mesic soils of medium textures (sandy loam to clay loams) but ranges into the lower reaches of frigid soils when other climatic variables are suitable. These soils are found most commonly in the order Ultisols and Alfisols. The best stands in the Sierra Nevada grow on deep, sandy loam soils developed from granitic rock. In the southern Cascade Range the best stands are on deep clay loams developed on basalt and rhyolite. In the Coast Range and Siskiyou Mountains in California and Oregon, the best stands are on soils derived from sandstone and shale.

Much of the terrain occupied by sugar pine is steep and rugged. Sugar pines are equally distributed on all aspects at lower elevations but grow best on warm exposures (southern and western) as elevation increases. Optimal growth occurs on gentle terrain at middle elevations.

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Special Uses

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Upper grades of old-growth sugar pine command premium prices for specialty uses where high dimensional stability, workability, and affinity for glue are essential. The wood is light (specific gravity, 0.34 ± 0.03) (3), resists shrinkage, warp, and twist, and is preferred for finely carved pattern stock for machinery and foundry casting. Uniformly soft, thin-celled spring and summer wood and straight grain account for the ease with which it cuts parallel to or across the grain, and for its satin-textured, lustrous finish when milled. Its easy working qualities favor it for molding, window and door frames, window sashes, doors, and other special products such as piano keys and organ pipes. Wood properties of young growth are not so well known. Pruning would undoubtedly be required to produce clear lumber during short rotations.

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Vegetative Reproduction

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Sugar pine does not sprout, but young trees can be rooted from cuttings. The degree of success is apparently under strong genetic control. In one trial the proportion of cuttings that rooted from different ortets from 3 to 6 years old ranged from 0 to 100 percent (27). As for most conifers, rootability diminishes rapidly with age of donor tree. Grafts, however, can be made from donors of all ages, with success rates from 70 to 80 percent common. Problems of incompatibility, frequent in some species such as Douglas-fir, are rare in sugar pine.

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Distribution

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Sugar pine extends from the west slope of the Cascade Range in north central Oregon to the Sierra San Pedro Martir in Baja California (approximate latitude 30° 30' to 45° 10' N.). Its distribution is almost continuous through the Klamath and Siskiyou Mountains and on west slopes of the Cascade Range and Sierra Nevada, but smaller and more disjunct populations are found in the Coast Ranges of southern Oregon and California, Transverse and Peninsula Ranges of southern California, and east of the Cascade and Sierra Nevada crests. Its southern extremity is an isolated population high on a plateau in the Sierra San Pedro Martir in Baja California. Over 80 percent of the growing stock is in California (49) where the most extensive and dense populations are found in mixed conifer forests on the west slope of the Sierra Nevada.

In elevation, sugar pine ranges from near sea level in the Coast Ranges to more than 3000 m (10,000 ft) in the Transverse Range. Elevational limits increase with decreasing latitude, with typical ranges as follows:

Cascade Range 335 to 1645 m (1,100 to 5,400 ft) Sierra Nevada 610 to 2285 m (2,000 to 7,500 ft) Transverse and Peninsula Ranges 1220 to 3000 m (4,000 to 10,000 ft) Sierra San Pedro Martir 2150 to 2775 m (7,065 to 9,100 ft)
- The native range of sugar pine.

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Brief Summary

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Pinaceae -- Pine family

Bohun B. Kinloch, Jr. and William H. Scheuner

Called "the most princely of the genus" by its discoverer, David Douglas, sugar pine (Pinus lambertiana) is the tallest and largest of all pines, commonly reaching heights of 53 to 61 m (175 to 200 ft) and d.b.h. of 91 to 152 cm (36 to 60 in). Old trees occasionally exceed 500 years and, among associated species, are second only to giant sequoia in volume. For products requiring large, clear pieces or high dimensional stability, sugar pine's soft, even-grained, satin-textured wood is unsurpassed in quality and value. The huge, asymmetrical branches high in the crowns of veteran trees, bent at their tips with long, pendulous cones, easily identify sugar pine, which "more than any other tree gives beauty and distinction to the Sierran forest" (25).

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Physical Description

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Tree, Evergreen, Monoecious, Habit erect, Trees without or rarely having knees, Tree with bark rough or scaly, Young shoots 3-dimensional, Buds resinous, Leaves needle-like, Leaves alternate, Needle-like leaf margins finely serrulate (use magnification or slide your finger along the leaf), Leaf apex acute, Leaves < 5 cm long, Leaves > 5 cm long, Leaves < 10 cm long, Leaves blue-green, Leaves white-striped, Needle-like leaves triangular, Needle-like leaves not twisted, Needle-like leaf habit erect, Needle-like leaves per fascicle mostly 5, Needle-like leaf sheath early deciduous, Twigs pubescent, Twigs viscid, Twigs not viscid, Twigs without peg-like projections or large fascicles after needles fall, Berry-like cones orange, Woody seed cones > 5 cm long, Seed cones bearing a scarlike umbo, Umbo with missing or very weak prickle, Umbo with obvious prickle, Bracts of seed cone included, Seeds brown, Seeds winged, Seeds unequally winged, Seed wings prominent, Seed wings equal to or broader than body.
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Pinus lambertiana

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Pinus lambertiana (commonly known as the sugar pine or sugar cone pine) is the tallest and most massive pine tree, and has the longest cones of any conifer. The species name lambertiana was given by the Scottish botanist David Douglas, who named the tree in honour of the English botanist, Aylmer Bourke Lambert. It is native to coastal and inland mountain areas along the Pacific coast of North America, as far north as Oregon and as far south as Baja California in Mexico.

Description

Growth

The sugar pine is the tallest and largest Pinus species, commonly growing to 40–60 meters (130–195 ft) tall, exceptionally to 82 m (269 ft) tall, with a trunk diameter of 1.2–2.5 m (3 ft 11 in – 8 ft 2 in), exceptionally 3.5 m (11 ft 6 in).[2] The tallest recorded specimen is 83.45 m (273 ft 9 in) tall, is located in Yosemite National Park, and was discovered in 2015.[3] The second tallest recorded was "Yosemite Giant", an 82.05 m (269 ft 2 in) tall specimen in Yosemite National Park, which died from a bark beetle attack in 2007. The tallest known living specimens today grow in southern Oregon and Yosemite National Park: one in Umpqua National Forest is 77.7 m (254 ft 11 in) tall and another in Siskiyou National Forest is 77.2 m (253 ft 3 in) tall. Yosemite National Park also has the third tallest, measured to 80.5 m (264 ft 1 in) tall as of June 2013; the Rim Fire affected this specimen, but it survived.

Old sugar pines in the Rogue River – Siskiyou National Forest, southern Oregon

The bark of Pinus lambertiana ranges from brown to purple in color and is 5–10 centimeters (2–4 in) thick.[2] The upper branches can reach out over 8 m (26 ft).[2] Like all members of the white pine group (Pinus subgenus Strobus), the leaves ("needles") grow in fascicles ("bundles") of five,[2] with a deciduous sheath. They are 5–11 cm (2–4+14 in) long.[4] Sugar pine is notable for having the longest cones of any conifer, mostly 10–50 cm (4–19+34 in) long,[2][5] exceptionally to 60 cm (23+12 in) long (although the cones of the Coulter pine are more massive); their unripe weight of 1–2 kilograms (2.2–4.4 lb) makes them perilous projectiles when chewed off by squirrels.[2] The seeds are 1–2 cm (1234 in) long, with a 2–3-centimeter (341+14-inch) long wing[5] that aids their dispersal by wind. Sugar pine never grows in pure stands, always in a mixed forest and is shade tolerant in youth.[6]

Distribution

The sugar pine occurs in the mountains of Oregon and California in the Western United States, and Baja California in northwestern Mexico; specifically the Cascade Range, Sierra Nevada, Coast Ranges, and Sierra San Pedro Martir. It is generally more abundant towards the south and can be found growing in elevations between 500 and 1,500 m (1,600 and 4,900 ft) above sea level.[2]

Genome

The massive 31 gigabase mega-genome of sugar pine has been sequenced in 2016 by the large PineRefSeq consortium.[7] This makes the genome one of the largest sequenced and assembled so far.[7]

The transposable elements that make up the megagenome are linked to the evolutionary change of the sugar pine. The sugar pine contains extended regions of non-coding DNA, most of which is derived from transposable elements. The genome of the sugar pine represents one extreme in all plants, with a stable diploid genome that is expanded by the proliferation of transposable elements, in contrast to the frequent polyploidization events in angiosperms.[8]

Almost ripe female cones

Embryonal growth

In late stage of embryonal development, the sugar pine embryo changes from a smooth and narrow paraboloid to a less symmetric structure. This configuration is caused by a transverse orientation of division planes in the upper portion of the embryo axis. The root initial zone is established, and the epicotyl develops as an anlage flanked by regions of that define the cotyledonary buttresses. At this stage, the embryo is composed of the suspensor, root initials and root cap region, hypocotyl-shoot axis, and the epicotyl. The upper (distal) portion of the embryo, which gives rise to the cotyledons and the epicotyl, is considered to be the shoot apex.[9]

Shoot apex

The apex has the following four zones:[10]

  1. The apical initials produce all cells of the shoot apex through cell division. It is located at the top of the meristem and the cells are larger in size compared to other cells on the surface layer.
  2. The central mother cell generates the rib meristem and the inner layers of the peripheral tissue zone through cell division. It presents a typical gymnosperm appearance and is characterized by cell expansion and unusual mitosis that occurs in the central region. The rate of mitosis increases on its outer edge.
  3. The peripheral tissue zone consists of two layers of cells that are characterized by dense cytoplasm and mitosis of high frequency.
  4. Lastly, the rib meristem is a regular arrangement of vertical files of cells which mature into the pith of the axis.

Etymology

Naturalist John Muir considered sugar pine to be the "king of the conifers". The common name comes from the sweet resin, which Native Americans used as a sweetener.[11] John Muir found it preferable to maple sugar.[12] It is also known as the great sugar pine. The scientific name was assigned by David Douglas, who was the first to describe it in 1826,[2] in honor of Aylmer Bourke Lambert.

Ecology

Wildlife

The large size and high nutritional value of the sugar pine seeds are appealing to many species. Yellow pine chipmunks (Neotamias amoenus) and Steller's jays (Cyanocitta stelleri) gather and hoard sugar pine seeds. Chipmunks gather wind-dispersed seeds from the ground and store them in large amounts. Jays collect seeds by pecking the cones with their beaks and catching the seeds as they fall out. Although wind is a main dispersion factor of sugar pine seeds, animals tend to collect and store them before the wind can blow them far.[13]

Black bears (Ursus americanus) rely on sugar pine seeds for their food source in the fall months within the Sierra Nevada. There is relationship between sugar pine seeds and oak acorns, as the bears will feed preferentially on those that are in a higher supply for that season. Both sugar pine and oak species are currently in decline, which can have a direct effect on black bear food sources within the Sierra Nevada.[14]

Threats

Sugar pine trees have been impacted by the invasive species of mountain pine beetles (Dendroctonus ponderosae) that are native to western North America. The beetles lay their eggs inside of the tree and inhibit the trees ability to defend itself against the invading species. The beetles also feed from the trees nutrients which slowly weakens the trees overall health, making the pines more susceptible to other threats like fires and fungal infections by white pine blister rust.[15] Blister rust can weaken the tree and enable further infestation by mountain pine beetles due to the lack of defense from the sugar pine.[16]

Sugar pine starting to succumb to white pine blister rust

The sugar pine has been severely affected by the white pine blister rust (Cronartium ribicola),[17] a fungus that was accidentally introduced from Europe in 1909. A high proportion of sugar pines have been killed by the blister rust, particularly in the northern part of the species' range that has experienced the rust for a longer period of time. The rust has also destroyed much of the Western white pine and whitebark pine throughout their ranges.[18] The U.S. Forest Service has a program (see link below) for developing rust-resistant sugar pine and western white pine. Seedlings of these trees have been introduced into the wild. The Sugar Pine Foundation in the Lake Tahoe Basin has been successful in finding resistant sugar pine seed trees and has demonstrated that it is important for the public to assist the U.S. Forest Service in restoring this species. However, blister rust is much less common in California, and sugar, Western white and whitebark pines still survive in great numbers there.[19]

The species is generally resistant to fire because of its thick bark and because it clears away competing species.[2] However, its mortality has been directly linked to dryer conditions and higher temperatures. Sugar pine trees grow in western North America, a region already impacted by climate change. Higher temperatures within a sugar pine forest can lower resin levels within the tree which will cause less protection against pathogens. At the same time the warmer winters make the survival of the pests and pathogens more likely. The weakened or dying trees then provide fuel to the forest fires, which may become more frequent and more intense, if the climate change results in warmer temperatures in summer, particularly if coupled with drier conditions and stronger winds.[20]

Protective efforts

Sugar pine trees are in a slow decline because of the several threats it faces: white pine blister rust, mountain pine beetles and climate change. Efforts to restore sugar pines and other white pine trees that have been impacted by invasive species, climate change and fires have been undertaken by governmental and non-governmental entities. One of the latter is a non-for-profit organization called Sugar Pine Foundation created in 2004 to plant sugar pine seeds in the Sierra Nevada along the border of California and Nevada.[19] They plant seedlings grown from seeds collected from blister rust resistant trees. In order to identify if the trees resistant to that pathogen, Sugar Pine Foundation tested over 500 sugar pine trees and have found 66 resistant trees.[19] The foundation is building a sugar pine population that is resistant to white pine rust because the fungus is a major threat and will continue to kill sugar pine trees at a very high rate.[21]

Uses

Bark of a sugar pine on Mount San Antonio

According to David Douglas, who was guided to the (exceptionally thick) tree specimen he was looking for by a Native American,[2] some tribes ate the sweetish seeds. These were eaten raw and roasted, and also used to make flour or pulverized into a spread.[2] Native Americans also ate the inner bark.[2] The sweet sap or pitch was consumed, in small quantities due to its laxative properties,[22] but could also be chewed as gum.[2] Its flavor is thought largely to be derived from the pinitol it contains.[2]

In the mid-19th century, the trees were used liberally as lumber during the California Gold Rush. In modern times they are used in much lower quantities, being spared for high-end products as with Western white pine.[2]

The odorless wood is also preferred for packing fruit, as well as storing drugs and other goods. Its straight grain also makes it a useful organ pipe material.[22]

Folklore

In the Achomawi creation myth, Annikadel, the creator, makes one of the 'First People' by intentionally dropping a sugar pine seed in a place where it can grow. One of the descendants in this ancestry is Sugarpine-Cone man, who has a handsome son named Ahsoballache.[23]

After Ahsoballache marries the daughter of To'kis the Chipmunk-woman, his grandfather insists that the new couple have a child. To this end, the grandfather breaks open a scale from a sugar pine cone, and secretly instructs Ahsoballache to immerse the scale's contents in spring water, then hide them inside a covered basket. Ahsoballache performs the tasks that night; at the next dawn, he and his wife discover the infant Edechewe near their bed.[23]

The Washo language has a word for sugar pine, simt'á:gɨm, and also a word for "sugar pine sugar", nanómba.

References

  1. ^ Farjon, A. (2013). "Pinus lambertiana". IUCN Red List of Threatened Species. 2013: e.T42374A2976106. doi:10.2305/IUCN.UK.2013-1.RLTS.T42374A2976106.en. Retrieved 13 November 2021.
  2. ^ a b c d e f g h i j k l m n o Arno, Stephen F.; Hammerly, Ramona P. (2020) [1977]. Northwest Trees: Identifying & Understanding the Region's Native Trees (field guide ed.). Seattle: Mountaineers Books. pp. 26, 30–35. ISBN 978-1-68051-329-5. OCLC 1141235469.
  3. ^ "3 Sierra sugar pines added to list of 6 biggest in world". Associated Press. South Lake Tahoe, California. 31 Jan 2021. Retrieved 13 Feb 2023.
  4. ^ Jepson Flora Project (ed.). "Pinus lambertiana". Jepson eFlora. The Jepson Herbarium, University of California, Berkeley.
  5. ^ a b Kral, Robert (1993). "Pinus lambertiana". In Flora of North America Editorial Committee (ed.). Flora of North America North of Mexico (FNA). Vol. 2. New York and Oxford – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  6. ^ Earle, Christopher J., ed. (2018). "Pinus lambertiana". The Gymnosperm Database.
  7. ^ a b Stevens, K.A.; et al. (2016). "Sequence of the Sugar Pine Megagenome". Genetics. 204 (4): 1613–1626. doi:10.1534/genetics.116.193227. PMC 5161289. PMID 27794028.
  8. ^ Gonzalez-Ibeas, Daniel; et al. (2016). "Assessing the Gene Content of the Megagenome: Sugar Pine (Pinus lambertiana)". G3 (Bethesda). 6 (12): 3787–3802. doi:10.1534/g3.116.032805. PMC 5144951. PMID 27799338.
  9. ^ Berlyn, Graeme P (1967). "The Structure of Germination in Pinus Lambertiana Dougl". Yale School of Forestry & Environmental Studies, Bulletin Series. 77.
  10. ^ Sacher, J.A. (1954). "Structure and Seasonal Activity of the Shoot Apices of Pinus Lambertiana and Pinus ponderosa". American Journal of Botany. 41 (9): 749–759. doi:10.1002/j.1537-2197.1954.tb14406.x.
  11. ^ "Sugar pine". Oregonencyclopedia.org. Retrieved 18 June 2017.
  12. ^ Saunders, Charles Francis (1976). Edible and Useful Wild Plants of the United States and Canada. Courier Dover Publications. p. 219. ISBN 0-486-23310-3.
  13. ^ Thayer, T; Vander Wall, S (2005). "Interactions between steller's jays and yellow pine chipmunks over scatter-hoarded sugar pine seeds". Journal of Animal Ecology. 74 (2): 365–374. doi:10.1111/j.1365-2656.2005.00932.x. JSTOR 3505625.
  14. ^ Mazur, R; Klimley, AP; Folger, K (2013). "Implications of the variable availability of seasonal foods on the home ranges of black bears, Ursus americanus, in the Sierra Nevada of California". Animal Biotelemetry. 1 (16): 16. doi:10.1186/2050-3385-1-16.
  15. ^ "Mountain pine beetle". Ontarios invading species awareness program. 2012. Archived from the original on 2020-09-27.
  16. ^ Van Mantgem, PJ; Stephenson, NL; Keifer, M; Keeley, J (2004). "Effects of an introduced pathogen and fire exclusion on the demography of sugar pine". Ecological Applications. 14 (5): 1590–1602. doi:10.1890/03-5109. JSTOR 4493673.
  17. ^ Moore, Gerry; Kershner, Bruce; Craig Tufts; Daniel Mathews; Gil Nelson; Spellenberg, Richard; Thieret, John W.; Terry Purinton; Block, Andrew (2008). National Wildlife Federation Field Guide to Trees of North America. New York: Sterling. p. 79. ISBN 978-1-4027-3875-3.
  18. ^ Maloney, P; Duriscoe, D; Smith, D; Burton, D; Davis, D; Pickett, J; Cousineau, R; Dunlap, J. "White Pine Blister Rust on High Elevation White Pines in California" (PDF). Archived from the original (PDF) on 2006-10-09. Retrieved 2007-02-05.
  19. ^ a b c "Sugar Pine Foundation". Sugarpinefoundation.org. Retrieved 18 June 2017.
  20. ^ Slack, A; Kane, J; Knapp, E; Sherriff, R (2017). "Contrasting impacts of climate and competition on large sugar pine growth and defense in a fire-excluded forest of the central sierra nevada". Forests. 8 (7): 244. doi:10.3390/f8070244.
  21. ^ Maloney, PE; Vogler, DR; Eckert, AJ; Jensen, CE; Neale, DB (2011). "Population biology of sugar pine (Pinus lambertiana Dougl.) with reference to historical disturbances in the Lake Tahoe Basin: Implications for restoration". Forest Ecology and Management. 262 (5): 770–779. doi:10.1016/j.foreco.2011.05.011.
  22. ^ a b Peattie, Donald Culross (1953). A Natural History of Western Trees. New York: Bonanza Books. p. 55.
  23. ^ a b Woiche, Istet (1992). Merriam, Clinton Hart (ed.). Annikadel: The History of the Universe as Told by the Achumawi Indians of California. Tucson: University of Arizona Press. ISBN 978-0-8165-1283-6. OCLC 631716557.
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Pinus lambertiana: Brief Summary

provided by wikipedia EN

Pinus lambertiana (commonly known as the sugar pine or sugar cone pine) is the tallest and most massive pine tree, and has the longest cones of any conifer. The species name lambertiana was given by the Scottish botanist David Douglas, who named the tree in honour of the English botanist, Aylmer Bourke Lambert. It is native to coastal and inland mountain areas along the Pacific coast of North America, as far north as Oregon and as far south as Baja California in Mexico.

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