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California Redwood

Sequoia sempervirens (D. Don) Endl.

Comments

provided by eFloras
Redwood is the only naturally occurring hexaploid conifer. It is one of only a few vegetatively reproducing conifers (from stump sprouts) and possibly the tallest tree species known. Winter buds, though small, are evident.

Redwood, including Sequoia sempervirens and Sequoiadendron giganteum , is the state tree of 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.
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Flora of North America Editorial Committee
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Description

provided by eFloras
Trees to 100(-110) m, suckering from base in native range; trunk buttressed at base, slightly tapered above, to 5 (-8) m d.b.h.; bark reddish brown or cinnamon colored, 15-25 cm thick, fibrous, exfoliating in broad, dark brown plates;

crown narrow; branches slender on young trees, finally stout, borne horizontally or basal ones deflexed. Leaves bright deep green adaxially, ca. 6 mm on main branchlets, 0.8-2 cm on lateral branchlets, midvein raised abaxially. Pollen cones ovoid, 1.5-2 mm; pollen yellow-green. Seed cones very small at pollination, maturing pale reddish brown, ovoid-elliptic or ovoid, 2-3.5 × 1.2-1.5 cm; cone scales shield-shaped, apically grooved, expanded into a rhomboid disc, occasionally with central mucro. Seeds pale brown, elliptic-oblong, ca. 1.5 mm; wing as wide as seed.

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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of China Vol. 4: 60 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Flora of China @ eFloras.org
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Wu Zhengyi, Peter H. Raven & Hong Deyuan
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Description

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Trees to ca. 110 m; trunk to 9 m diam.; crown conic and monopodial when young, narrowed conic in age. Bark reddish brown, to ca. 35 cm thick, fibrous, ridged and furrowed. Branches downward sweeping to slightly ascending. Leaves 1--30 mm, generally with stomates on both surfaces, the free portion to 30 mm, those on leaders, ascending branchlets, and fertile shoots divergent to strongly appressed, short-lanceolate to deltate, those on horizontally spreading to drooping branchlets mostly linear to linear-lanceolate, divergent and in 2 ranks, with 2 prominent, white abaxial stomatal bands. Pollen cones nearly globose to ovoid, 2--5 mm, borne singly on short terminal or axillary stalks. Seed cones 1.3--3.5 cm. Seeds flattened, 3--6 mm, leathery. 2 n = 66.
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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
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eFloras.org
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Distribution

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Calif., Oreg.
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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.
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Flora of North America @ eFloras.org
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Flora of North America Editorial Committee
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Habitat

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Coastal redwood forests; generally below 300 m, occasionally to 1000m.
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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
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eFloras.org
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Habitat & Distribution

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Cultivated. Fujian, Guangxi, Jiangsu (Nanjing Shi), Jiangxi, Taiwan, Zhejiang (Hangzhou Shi) [native to W United States].
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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 China Vol. 4: 60 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
source
Flora of China @ eFloras.org
editor
Wu Zhengyi, Peter H. Raven & Hong Deyuan
project
eFloras.org
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eFloras

Synonym

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Taxodium sempervirens D. Don in Lambert, Descr. Pinus 2: [24]. 1824; Sequoia gigantea Endlicher (1847), not (Lindley) Decaisne (1854).
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copyright
Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of China Vol. 4: 60 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
source
Flora of China @ eFloras.org
editor
Wu Zhengyi, Peter H. Raven & Hong Deyuan
project
eFloras.org
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eFloras

Synonym

provided by eFloras
Taxodium sempervirens D. Don in Lambert, Descr. Pinus 2: 24. 1824
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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
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Brief Summary

provided by EOL authors
Redwood is a native, evergreen, long-lived (greater than 2,200 years), monoecious tree [38,40] (monoecious = "having reproductive organs typical of both sexes in a single individual"). Redwoods are among the world's tallest trees; trees over 200 feet (61 m) are common, and many are over 300 feet (91 m) [40]. The largest tree thus far was measured at 364 feet (110.3 m) in height and 20 feet (6.1 m) in d.b.h. ("diameter at breast height") [44]. The root system is composed of deep, widespreading lateral roots with no taproot [40,44]. The bark is up to 12 inches (30 cm) thick and quite fibrous [44]. Redwood self-prunes well in dense stands [40]; the base of the bole is strongly buttressed [38]. Redwood is endemic to the coastal area of northern California and southwestern Oregon. The redwoods occupy a narrow strip of land approximately 450 miles (724 km) in length and 5 to 35 miles (8-56 km) in width. The northern boundary of its range is marked by two groves on the Chetco River in the Siskiyou Mountains within 15 miles (25 km) of the California-Oregon border [22,40]. The southern boundary of redwood's range is marked by a grove in Salmon Creek Canyon in the Santa Lucia Mountains of southern Monterey County, California [40]. Redwood occurs in a maritime Mediterranean climate, where the winters are cool and rainy, and the summers are dry. The mean precipitation is 70 inches (180 cm), with 90 percent falling between October and May. The dry summers are mitigated by a heavy fog belt [30]. The fog reduces the drought stress of this hydrophilic plant by reducing evapotranspiration and adding soil moisture. Redwoods beyond the fog belt appear to be limited to areas of high moisture. Currently there is considerable debate over the link between the fog belt and redwood distribution [11].

The Man Who Planted Trees

provided by EOL authors

Great read, easy to reference, and Mr. Jim Robins does an incredible job of describing many other tree species as well.

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Broad-scale Impacts of Fire

provided by Fire Effects Information System Plants
More info for the terms: top-kill, tree

Young trees originating from stump sprouts have a higher rate of
top-kill after fire than those originating from seedlings (see fire case
study) [16].

Basal wounding provides a vector for heart rot to enter the tree.  Once
this has occurred, recurring fires and basal decay produce large basal
cavities, called goosepens, that weaken the tree [40].
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bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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/

Broad-scale Impacts of Plant Response to Fire

provided by Fire Effects Information System Plants
More info for the terms: prescribed fire, root crown, severity, top-kill, tree

After crown-kill redwood sprouts new foliage from dormant buds along the
bole.  The bole is covered with fine feathery foliage extending 2 to 3
feet (0.6-0.9 m) out from the bole.  This manifestation is called a
fire-column.  Over time the narrowed crown will again develop into a
typical crown.  During the first 4 postfire years the tree will produce
very few strobili [40].

After top-kill, the number of sprouts per root crown depends on the
severity of the fire.  Severe heat influx to the root crown kills more
of the dormant buds, thus reducing the number of sprouts; however, this
allocates more of the carbohydrate reserves to fewer sprouts, which
results in larger and taller sprouts [14].

In northwestern California, Finney and Martin [14,16] found stump sprouts
were less likely to survive prescribed fire than redwood seedlings.  Large
redwoods survived prescribed fire.  For further information, see Fire Case Studies.
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cc-publicdomain
bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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
redwood
California redwood
coast redwood
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bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

provided by Fire Effects Information System Plants
More info for the terms: cover, tree

Redwood forests provide hiding and thermal cover for Roosevelt elk,
black-tailed deer, and a variety of small mammals [24,45,48,50].

The pileated woodpecker generally selects broken tree tops or snags with
rot for nesting cover.  The softness of redwood, however, allows the
pileated woodpecker to use green trees of adequate size.  In one study
only half the nests of pileated woodpeckers were in redwoods that had
broken tops with rot, while the other half were in sound green trees
with no sign of decay in the excavation chips [25].

In California, the state-endangered marbled murrelet nests exclusively
in coastal old-growth redwood forests [46].
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cc-publicdomain
bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

provided by Fire Effects Information System Plants
More info for the terms: monoecious, tree

Redwood is a native, evergreen, long-lived (greater than 2,200 years),
monoecious tree [38,40].  Redwoods are among the world's tallest trees;
trees over 200 feet (61 m) are common, and many are over 300 feet (91 m)
[40].  The largest tree thus far was measured at 364 feet (110.3 m) in
height and 20 feet (6.1 m) in d.b.h. [44].  The root system is composed
of deep, widespreading lateral roots with no taproot [40,44].  The bark
is up to 12 inches (30 cm) thick and quite fibrous [44].  Redwood
self-prunes well in dense stands [40]; the base of the bole is strongly
buttressed [38].
license
cc-publicdomain
bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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
Redwood is endemic to the coastal area of northern California and
southwestern Oregon.  The redwoods occupy a narrow strip of land
approximately 450 miles (724 km) in length and 5 to 35 miles (8-56 km)
in width.  The northern boundary of its range is marked by two groves on
the Chetco River in the Siskiyou Mountains within 15 miles (25 km) of
the California-Oregon border [22,40].  The southern boundary of
redwood's range is marked by a grove in Salmon Creek Canyon in the Santa
Lucia Mountains of southern Monterey County, California [40].
license
cc-publicdomain
bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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 terms: fire interval, fire regime, forest, mean fire interval, root crown

Fire has had an ecological role in the redwood forest type [53].  The
mean fire interval (MFI) prior to human occupation was approximately 135
to 350 years, and after human influx (about 11,000 years ago) decreased
to approximately 17 to 82 years [21].  Redwood has adapted to this fire
regime, and mature redwoods are considered very resilient to fire.  The
thick bark; great height; and ability to sprout from the root crown or
from dormant buds located under the bark of the bole and branches are
adaptations that allow redwood to survive cool to hot fires [16].

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".
license
cc-publicdomain
bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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
More info for the terms: duff, fire exclusion, fire interval, fire management, fire regime, fireline intensity, flame length, forest, frequency, fuel, fuel loading, fuel moisture, litter, natural, prescribed fire, reburn, seed, top-kill

A fire regime where prescribed fire substitutes for lightning and
now-absent aboriginal ignitions may have to be implemented to maintain
or reestablish presettlement conditions in old-growth or cutover redwood
forests [15].  McBride and others [34] recommend that both frequency
distributions of fire intervals and an analysis of the pattern of fire
intervals be used as a basis for determining reburn intervals for
prescribed fire.  They evaluated the fire history of redwood forest
stands in Muir Woods National Monument and, because of the highly skewed
frequency distribution observed in this type, suggested that the average
fire interval would be inappropriate to use as a reburn interval.
Instead a combination of shorter than the average and longer than the
average natural fire interval was recommended.  In areas where fire has
been excluded for many decades, a prescribed fire program should start
with two short-interval fires (less than average interval) to reduce
high fuel accumulations.  Once the fuel load has been reduced, a burning
pattern of two short fire intervals followed by a long interval should
be implemented [34].

Person and Hallin [43] reported that regeneration was 5 to 10 times
greater on cuts with moderate to hot slash fires than on those with cool
or no slash fires.  Hallin [23] proposed the following guidelines for
slash fires:

       (1)  burn at night
       (2)  do not burn during the dry season (June thru September)
       (3)  light winds
       (4)  keep the area small (less than 40
acres [16 ha])
       (5)  slash loads pulled away from advance regeneration

If sprouts are to be used as part of stand regeneration, the stumps
should not be debarked or severely burned during slash disposal, as
these actions will result in lowered sprout stocking [10].

Finney [14] has developed equations to estimate the fuel loading of the
forest floor in redwood stands based on forest floor depth.

FIRE CASE STUDY
SPECIES: Sequoia sempervirens
FIRE CASE STUDY CITATION :
Griffith, Randy Scott., compiler. 1992. Prescribed underburning in young-growth
redwood forests. In: Sequoia sempervirens. In: Fire Effects Information
System, [Online]. U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences Laboratory (Producer).
Available: https://www.fs.fed.us
/database/feis/ [
var months = new Array(12);
months[0] = "January";
months[1] = "February";
months[2] = "March";
months[3] = "April";
months[4] = "May";
months[5] = "June";
months[6] = "July";
months[7] = "August";
months[8] = "September";
months[9] = "October";
months[10] = "November";
months[11] = "December";
var date = new Date();
var year = date.getFullYear();
var month = date.getMonth();
var day = date.getDate();
document.write(year+", "+months[month]+" "+day);
].


REFERENCES :
Finney, Mark Arnold. 1991. Ecological effects of prescribed and
simulated fire on the coast redwood (Sequoia sempervirens (D. Don)
Endl.). Berkeley, CA: University of California. 179 p. Dissertation.
[14].

Finney, M. A.; Martin, R. E. 1991. Prescribed underburning and some
initial effects in young-growth coast redwood forests of California.
In: Andrews, Patricia L.; Potts, Donald F., eds. Proceedings, 11th
annual conference on fire and forest meteorology; 1991 April 16-19;
Missoula, MT. SAF Publication 91-04. Bethesda, MD: Society of American
Foresters: 328-334. [16].


SEASON/SEVERITY CLASSIFICATION :
May     1989  Low consumption burn
June    1990  Low consumption burn
October 1989  High consumption burn


STUDY LOCATION :
The study took place in two of California's state parks, Annadel State
Park and Humboldt Redwoods State Park.  Each area had 16 plots.  Annadel
State Park is located approximately 5 kilometers (3 mi) east of Santa
Rosa, California.  The coordinates are 38 degrees 25 minutes North
latitude, 122 degrees 35 minutes West longitude, and the legal
description is T7N R7W SW1/4 of section 25.

Humboldt State Park is located approximately 320 kilometers (192 mi)
north of Annadel State Park near Weott, California.  The coordinates are
39 degrees 16 minutes North latitude, 123 degrees 45 minutes West
longitude, and the legal description is T1S R2E S1/2 NW1/4 of section
20.


PREFIRE VEGETATIVE COMMUNITY :
At the Annadel site the forest was young growth with diameters between 5
and 40 centimeters (2-16 in).  The age of the stand was between 120 and
140 years.  Evidence on the site suggests that the stand developed from
sprouts after fire exclusion in brush fields began in the mid-1800s.
The fire regime in the area prior to exclusion was every 2 to 6 years.

The Humboldt site was also a young growth forest with diameters between
5 and 40 centimeters (2-16 in).  This stand developed from natural
regeneration after logging approximately 60 to 80 years prior to the
study.  The past fire regime in the Humboldt area was every 5 to 25
years.


TARGET SPECIES PHENOLOGICAL STATE :
The prescribed burns took place when the trees were actively growing;
the burn in October occurred at the onset of seed dispersal.


SITE DESCRIPTION :
On the Annadel site the plots were located on a northern aspect with a
slopes ranging from 30 to 40 percent.  The elevation of the plots ranged
from 240 to 350 meters (792-1,155 ft).

On the Humboldt site the plots were located on a southern exposure with
slopes ranging from 10 to 40 percent.  The elevation of the plots ranged
from 350 to 450 meters (1,155-1,485 ft).


FIRE DESCRIPTION :
To achieve different burn severities on the plots a variety of firing
techniques were used.  To achieve low fireline intensities, backing and
flanking fires were used, and to obtain higher fireline intensities,
strip head fires were used, with the strips ranging from 3 to 6 meters
(10-20 ft) wide to regulate fuel availability and build-up of fireline
intensity.

On the Annadel site, two plots were burned on May 17, 1989; one plot on
May 25, 1989; five plots were burned on May 30, 1989; and 8 plots were
burned on October 17, 1989.  Conditions were as follows:

                        Annadel Site

Date          Air           Rel.       Wind
              Temp.         Humd.      Speed
              (C)           (%)        (m/s)
 
5/17/89       16-18         55-70       0-2.2
5/25/89       18            38-45       0-1.3
5/30/89       18-21         40-45       0-2.2
10/17/89      17-24         35-52       0

On the Humboldt site, seven plots were burned on October 10, 1989; one
plot was burned on October 11, 1989; and eight plots were burned on June
27, 1990.  Conditions were as follows:

                         Humboldt Site

Date          Air           Rel.        Wind
              Temp.         Humd.       Speed
              (C)           (%)         (m/s)

10/10/89      13-24         48-75       0-2.4
10/11/89      16-22         60-72       0-1.3
6/27/90       20            56          0-1.3

For more specific information on burn conditions and ignition pattern
for each of the 16 plots on the two sites, see Finney 1990.

The fuel loading on both sites ranged from 29 to 55 tonnes per hectare
(32-61 tn/a) of litter and duff.  The small woody fuels (0-7.62 cm in
diameter) ranged from 9 to 20 tonnes per hectare with the large woody
fuels (> 7.62 cm) being highly variable.

The flame length ranged from 0.27 to 2.07 meters (0.9-6.8 ft) which
relates to a fireline intensity of 40 to 1833 kilowatts per meter
second.  

The fuel consumption varied from 15 to 68 tonnes per hectare.  The
percentage of fuel consumed ranged from 23 to 100 percent.  Fuel
consumption was found to be positively related to fuel loading.

Finney provides in-depth information on these parameters.


FIRE EFFECTS ON TARGET SPECIES :
Most redwoods greater than 15 to 20 centimeters d.b.h. survived the most
severe prescribed fire with 100 percent surface fuel consumption and 100
percent crown scorch.

Redwoods originating from stump sprouts after logging were found to have
higher rates of top-kill than those originating from seedlings.  This
was linked to higher fuel concentrations (2 to 5 times that between
clumps), the stump, and heat convection currents drafting inward and
around the circular clump of trees.

As redwoods achieve greater d.b.h. the probability of top-kill
decreases.


FIRE MANAGEMENT IMPLICATIONS :
Flame length and fuel consumption were found to be the most important
parameters in determining top-kill and basal sprouting.  These
parameters can be easily controlled by use of different firing patterns
and fuel moisture to achieve the desired effects from a prescribed fire.

A regime of periodic prescribed fire would elevate the probability of
sprout regeneration being top-killed by preventing large fuel
accumulations.
license
cc-publicdomain
bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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 Implications

provided by Fire Effects Information System Plants
More info for the terms: fuel, fuel moisture, prescribed fire, top-kill

Flame length and fuel consumption were found to be the most important
parameters in determining top-kill and basal sprouting.  These
parameters can be easily controlled by use of different firing patterns
and fuel moisture to achieve the desired effects from a prescribed fire.

A regime of periodic prescribed fire would elevate the probability of
sprout regeneration being top-killed by preventing large fuel
accumulations.

Growth Form (according to Raunkiær Life-form classification)

provided by Fire Effects Information System Plants
More info on this topic.

More info for the term: phanerophyte

   Phanerophyte (megaphanerophyte)
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bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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 terms: grassland, vine

Redwood occurs in a maritime Mediterranean climate, where the winters
are cool and rainy, and the summers are dry.  The mean precipitation is
70 inches (180 cm), with 90 percent falling between October and May.
The dry summers are mitigated by a heavy fog belt [30].  The fog reduces
the drought stress of this hydrophilic plant by reducing
evapotranspiration and adding soil moisture.  Redwoods beyond the fog
belt appear to be limited to areas of high moisture.  Currently there is
considerable debate over the link between the fog belt and redwood
distribution [11].

Preferred sites for redwood stands are alluvial fans, coastal plains,
and benches along large streams [40].  The size of a redwood can be site
dependent:  a 400-year-old specimen on a hillside had a d.b.h. of 2 feet
(0.6 m), while a 600-year-old specimen on an alluvial fan had a d.b.h.
of 12 feet (3.6 m) [4].

Elevation:  Redwood occurs at elevations ranging from sea level to 3,000
feet (0-915 m), but most stands occur from 100 to 2,320 feet (100-703 m)
[11,40].  Redwoods are sensitive to salt spray [40], and are usually
separated from the coast by intervening grassland [22]

Soils:  Redwood has a strong affinity for deep, moist soils in the
Inceptisol and Ultisol soil orders [40].  The common parent materials
are graywacke sandstones, shales, and conglomerates [30].

Associates:  In addition to those previously listed under Distribution
and Occurrence, overstory associates include Sitka spruce (Picea
sitchensis), Pacific yew (Taxus brevifolia), California torreya (Torreya
californica), Gowen cypress (Cupressus goveniana), bishop pine (Pinus
muricata), Monterey pine (P. radiata), bigleaf maple (Acer macrophyllum),
Oregon white oak (Quercus garryana), and Oregon ash (Fraxinus latifolia)
[40].

Understory associates include vine maple (Acer circenatum), chittam bark
(Rhamnus purshiana), evergreen huckleberry (Vaccinium ovatum), Pacific
rhododendron (Rhododendron macrophyllum), salmon berry (Rubus
spectabilis), and evergreen ceanothus (Ceanothus velutinus) [40].
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bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

provided by Fire Effects Information System Plants
More info on this topic.

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

   229  Pacific Douglas-fir
   231  Port-Orford-cedar
   232  Redwood
   234  Douglas-fir - tanoak - Pacific madrone
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bibliographic citation
Griffith, Randy Scott. 1992. Sequoia sempervirens. 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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More info on this topic.

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
   FRES27  Redwood
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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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This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the term: forest

   K002  Cedar - hemlock - Douglas-fir forest
   K006  Redwood forest
   K028  Mosaic of K002 and K026
   K029  California mixed evergreen forest
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

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More info for the terms: top-kill, tree

The effect of fire on redwood varies depending on the size of the tree.
The bark of young trees (less than 8 inches [20 cm] d.b.h.) is generally
too thin to protect the cambium from damage, and trees of this size are
usually top-killed by cool to hot fires [16].  The thick bark of mature
redwood insulates the cambium from the heat of the fire [15], and in
many cases, fire may only reduce bark thickness [40].  Under more severe
circumstances, such as stand-replacing fires, basal wounding and
top-kill occurs [40].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

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Redwood forests provide habitat for variety of mammals, aviafauna,
reptiles, and amphibians [7,45,48].  Remnant old-growth redwood stands
provide habitat for the federally threatened spotted owl and the
California-endangered marbled murrelet [1,46].

In settlement times fire scar cavities at the base of larger redwood
boles were used as goose pens; hence the name "goosepens" has been used
to denote fire scar cavities [14].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

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More info for the terms: codominant, forest, natural, series

Redwood is listed as a dominant or codominant overstory species in the
following publications:

Coast redwood ecological types of southern Monterey County, California [11].
Terrestrial natural communities of California [26].
The redwood forest and associated north coast forests [58].
Forest associations of Little Lost Man Creek, Humboldt County,
  California: Reference-level in the hierarchical structure of
  old-growth coastal redwood vegetation [30].
Preliminary plant associations of the Siskiyou Mountain Province [5].
Tanoak series of the Siskiyou Region of southwest Oregon [6].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

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More info for the term: tree

Tree
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

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More info for the terms: competition, forest, natural, selection, shrubs

Wildlife:  The marbled murrelet is dependent on old-growth redwood
forests for nesting habitat.  This bird is listed as endangered in
California and is under consideration for federal protection as a
threatened species in California, Oregon, and Washington [1].
Old-growth redwood forests of northern California also provide critical
habitat for the federally endangered northern spotted owl [1].

Black-tailed deer numbers increase after clearcutting in the redwood
forest type as a result of the sudden increase in available understory
forage.  After canopy closure (20 to 30 years), black-tailed deer
numbers decrease rapidly [50].

        Years after clearcut    Number of deer

         0 to 5                      43
         5 to 10                    142
        10 to 15                     21
        15 to 20                     21
        20 to 25                      8
        25 to 30                      8     


Competition:  Evergreen hardwoods are strong competitors in the redwood
forest type.  Tanoak (Lithocarpus densiflorus) and Pacific madrone
(Arbutus menziesii) often sprout when cut, and reoccupy the site
before redwood.  These competitors can be controlled by trunk injections
of triclopyr (Garlon 3A), with two to three treatments over a 4- to
5-year period giving the best results.  Foliar spraying with triclopyr
can also control hardwoods but has adverse effects on redwood [56].

Mulching and the use of ground covers increase survival of planted
seedlings by reducing water evaporation and reducing competition from
shrubs [35].  Seedling survival can also be enhanced with the use of
shades [2].

Damage:  Damaging agents include insects, branch canker (Coryneum spp.),
and heart rots (Poria sequoiae, P. albipellucida).  The insects
associated with redwood cause no significant damage, but the branch
canker girdles stems and branches, which can be especially harmful in
plantations.  Heart rots cause extensive cull in the redwood forest type
[40].

Wood rats girdle and strip the bark of redwood seedlings, and can
seriously limit redwood regeneration.  Where this is a problem, site
preparation should include destroying wood rat nesting areas [49].

Redwood is susceptible to damage from soil compaction in areas of heavy
foot traffic [4].

Silviculture:  The preferred silvicultural system for harvesting
redwoods is small clearcuts (30 to 40 acres) [10,41].  Boe [9] provides
information on the three silvicultural systems used in the redwood
forest type:  clearcut, shelterwood, and selection cut.

Other:  Namkoong and Roberds [39] developed an extinction model for
redwood.  Their findings reveal there is a small probability of
extinction due to natural processes, which can easily be circumvented by
planting.
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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  OR  HI
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Other uses and values

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The cultivars 'Nana Pendula' and 'Prostrata' are grown extensively as
ornamentals due to their reduced size [28].  Redwood has been planted in
New Zealand, Australia, and Europe [40].

Native Americans used redwood in the construction of canoes and as grave
markers [51].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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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More info for the term: seed

Redwood female strobili become receptive and pollen is shed from late
November to early March.  Female strobili start ripening in September of
the first year.  Mature female strobili can be identified when their
color changes from green to greenish yellow.  Seed dispersal begins in
late October, with most of the seeds being dispersed from November to
February [8].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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/

Plant Response to Fire

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More info for the terms: root crown, seed, top-kill

After fires that destroy all aboveground portions, both mature and young
redwoods will sprout from the root crown [40]; even seedlings have the
ability to sprout after top-kill [30].  After fires that destroy the
crown, redwoods greater than 8 inches (20 cm) will sprout from numerous
dormant buds along the bole and produce new foliage (see fire case
study) [14,16,40].

Redwood can also reestablish after fire via on-site and off-site seed
[43].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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 terms: crown residual colonizer, ground residual colonizer, secondary colonizer, seed, tree

   Tree with adventitious-bud root crown/root sucker
   Ground residual colonizer (on-site, initial community)
   Crown residual colonizer (on-site, initial community)
   Secondary colonizer - on-site seed
   Secondary colonizer - off-site seed
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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

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More info for the terms: adventitious, density, epigeal, formation, layering, root crown, seed, top-kill, tree

Redwood reproduces both sexually and asexually.  The male and female
strobili are borne separately on different branches.  Redwood begins
producing seeds at 5 to 15 years of age.  Large seed crops occur
frequently, but viability of the seed is low [8].  A dry period during
pollination allows better pollen dispersal and improves seed viability.
The seeds are small and light, averaging 120,000 seeds per pound
(265,000 seeds/kg).  The wings are not effective for wide dispersal
[19], and seeds are dispersed by wind an average of only 200 to 400 feet
(61-122 m) from the parent tree [40].

Redwood seeds do not require pretreatment to germinate.  Germination is
epigeal [40]; the best seedbed is moist mineral soil with some shade
[17,36].  Germination rates are generally low due to low viability
rather than to dormancy.  Germination rates with a mean of 10 percent
are the norm [8].

Seedlings require adequate moisture to survive.  The roots of redwood
seedlings do not have root hairs and are thus inefficient at extracting
soil moisture.  Once established seedlings can obtain remarkable growth
rates in the first season.  Growth of 18 inches (46 cm) is not uncommon.
Older saplings (4 to 10 years old) can grow 6.5 feet (2.0 m) in one
growing season [40].

Redwoods can reproduce asexually by layering or sprouting from the root
crown or stump.  Sprouts from the root crown are generally favored for
tree crops [10]; sprouts originating from the stump are generally not as
vigorous as root-crown sprouts, and are very susceptible to wind throw
[40].  Sprouts originate from dormant or adventitious buds at or under
the surface of the bark [17,40].  The formation of these buds occurs at
a young age, as even seedlings have been observed to sprout after
top-kill [30].  The sprouting capacity of redwood decreases with size
and age [17].  Sprouting appears to be the greatest on the downhill side
of the tree [14].  Within a short period after sprouting each sprout
will develop its own root system, with the dominant sprouts forming a
ring of trees around the parent root crown [40].  The mean crop tree
sprouting potential per root crown is five, which adds many crop trees
to a given site [10].

Sprouts can achieve heights of 7 feet (2.1 m) in a single growing
season.  Shading does not decrease sprout height, but it does reduce the
number and weight of sprouts [14].  Density of sprouts also affects
sprout vigor; the higher the density, the less vigorous the sprouts
[40].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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/

Season/Severity Classification

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May     1989  Low consumption burn
June    1990  Low consumption burn
October 1989  High consumption burn

Site Description

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On the Annadel site the plots were located on a northern aspect with a
slopes ranging from 30 to 40 percent.  The elevation of the plots ranged
from 240 to 350 meters (792-1,155 ft).

On the Humboldt site the plots were located on a southern exposure with
slopes ranging from 10 to 40 percent.  The elevation of the plots ranged
from 350 to 450 meters (1,155-1,485 ft).

Successional Status

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More info for the terms: climax, tree

Facultative Seral Species
Obligate Climax Species

Redwood is classified as a shade-tolerant to very shade-tolerant species
due to its high photosynthetic capacity at low light levels [40].
Redwood releases well even at quite an old age.  One specimen after
1,000 years released from 30 to 6 rings per inch (12-2.4 rings/cm) [19].

There is some debate over the classification of redwood as a climax
species.  Some consider redwood a climax species, while others consider
it a fire-dependent seral species [15,41,54,55].  Osburn and Lowell [41]
reported that if fire is excluded from Redwood National Park over the
next 2,000 years redwood will disappear, and Sitka spruce, western
hemlock (Tsuga heterphylla), and western redcedar (Thuja plicata) will
dominate.  Viers [55] on the other hand reported that redwood is a
climax species in the vicinity of Redwood National Park because it
maintains uneven age distributions with or without fire.

After disturbance redwood dominates in early seres due to its ability to
sprout [27,58].

In the floodplain environment redwood deploys what has been called "the
endurer strategy."  After flooding and stem burial, redwood will develop
a new and higher lateral root system from buried buds on the bole of the
tree.  While the repeated flooding and deposition of soil (often to
depths of 30 inches [76 cm]) kills competing vegetation, redwood endures
[3,40,58].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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/

Synonyms

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Taxodium sempervirens (D. Don) Lamb.
Steinhauera sempervirens (Voss. S.) Presl.
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Taxonomy

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The currently accepted scientific name of redwood is Sequoia
sempervirens (D. Don) Endl. Redwood is a member of the Taxodium
(Taxodiaceae) family [44]. There are no recognized subspecies,
varieties, or forms.
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Value for rehabilitation of disturbed sites

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More info for the terms: natural, restoration, seed, tree

In a large cutover area acquired by Redwood National Park, both
plantings and natural colonization of redwood on outsloped (recontoured
into the hillside) logging roads were used with good success.  This
treatment curtailed erosion in the park by an estimated 6.6 million
cubic feet (0.2 mil m3) [33].

Redwood was one of a number of native species used successfully to
reclaim a riparian ecosystem in a city park in Berkeley; redwoods on the
site had a high survival rate [57].

Redwood can be propagated via seed or cuttings.  Seeds should be sown
from December to April.  If planting with a seed drill, the recommended
depth is 0.125 inch (0.32 cm), with a seeding rate that will yield 30
seedlings per square foot (333 seedlings/sq m) [8].  Cuttings from 2- to
3-year-old seedlings produce the highest percentage of rooted cuttings
(up to 90 percent); cuttings from older trees are more difficult to root
[36,40].  Hedging (close-cropping) can maintain the rooting capabilities
of the donor tree.  By repeated hedging a single donor seedling and its
clones can produce a million cuttings in 3 years [40].  Redwood can also
be successfully propagated in plant tissue culture.  The callus can be
induced to generate cultured plantlets.  The cultured plantlets are
usually twice the size of seedlings the same age [40].

Millar and Libby [37] have developed guidelines for redwood seed
collection and for the use of redwood in the restoration of disturbed
areas.
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Wood Products Value

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Redwood is one of California's most valuable timber species [36].  The
wood is soft, weak, easily split, and very resistant to decay
[38,40,44].  The clear wood is used for dimension stock and shingles
[44].  Redwood burls are used in the production of table tops, veneers,
and turned goods [40].
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Griffith, Randy Scott. 1992. Sequoia sempervirens. 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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Redwood is a principal species in only one forest cover type, Redwood (Society of American Foresters Type 232) (42), but is found in three other Pacific Coast types, Pacific Douglas-Fir (Type 229), Port-Orford-Cedar (Type 231), and Douglas-Fir-Tanoak-Pacific Madrone (Type 234).

Pure stands of redwood are found only on some of the best sites, usually the moist river flats and gentle slopes below 305 m (1,000 ft). Although redwood is a dominant tree throughout its range, generally it is mixed with other conifers and broad-leaf trees.

Douglas-fir (Pseudotsuga menziesii) is well distributed throughout most of the redwood type. Distributions of other conifer associates are more limited. Significant species on the coastal side of the redwood type are grand fir (Abies grandis) and western hemlock (Tsuga heterophylla) north from northern Sonoma County, CA, and Sitka spruce (Picea sitchensis) north from the vicinity of Humboldt Bay, CA.

Conifers associated less commonly on the coastal side of the redwood type are Port-Orford-cedar (Chamaecyparis lawsoniana), Pacific yew (Taxus brevifolia), western redcedar (Thuja plicata), and California torreya (Torreya californica). Other conifers found with redwood include Gowen cypress (Cupressus goveniana) and several species of pine, including bishop pine (Pinus muricata), knobcone pine (P. attenuata), lodgepole pine (P. contorta), Monterey pine (P. radiata), and sugar pine (P. lambertiana).

The two hardwoods most abundant and generally distributed in the redwood region are tanoak (Lithocarpus densiflorus) and Pacific madrone (Arbutus menziesii). Other hardwoods found with redwood include vine maple (Acer circinatum), bigleaf maple (A. macrophyllum), red alder (Alnus rubra), giant chinkapin (Castanopsis chrysophylla), Oregon ash (Fraxinus latifolia), Pacific bayberry (Myrica californica), Oregon white oak (Quercus garryana), cascara buckthorn (Rhamnus purshiana), willows (Salix spp.), and California-laurel (Umbellularia californica).

Of the great variety of lesser vegetation found in association with redwood, these species are especially common: bracken (Pteridium aquilinum var. lanuginosum), sword fern (Polystichum munitum), salal (Gaultheria shallon), blueblossom (Ceanothus thyrsiflorus), California huckleberry (Vaccinium ovatum), Pacific rhododendron (Rhododendron macrophyllum), salmonberry (Rubus spectabilis), coyote-brush (Baccharis pilularis), and snowbrush (Ceanothus velutinus).

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Climate

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The mild climate of the redwood forest region can be classed broadly as super-humid or humid. Mean annual temperatures vary between 10° and 16° C (50° and 60° F). Differences between mean annual maximum and mean annual minimum temperatures vary from -12° C (10° F) for coastal points to -1° C (30° F) for the eastern edge of the redwood type. Temperatures rarely drop below -9° C (15° F) or rise above 38° C (100° F). The frost-free period varies from 6 to 11 months (34).

Annual precipitation varies between 640 and 3100 mm (25 and 122 in) and is mostly winter rain, although snow sometimes covers the highest ridges. Generally, January is the wettest month and July is the driest. With substantial precipitation in all months except summer, only slight summer drought on deep soils, and mild winters, the climate is productive, and some of the world's grandest forests are indigenous to it (34).

The frequent summer fogs that blanket the redwood region seem to be more significant than the amount of precipitation in delineating the redwood type. The major effect of fog is to decrease water loss from evaporation and transpiration. An additional effect of condensation and fog drip from tree crowns is an increased soil moisture supply during the dry summers (1). The natural range of redwood is limited to areas where heavy summer fogs from the ocean provide a humid atmosphere, although its successful growth in plantations or amenity plantings is not as limited. Redwood is among the most successful trees in the Central Valley of California, and at low elevations in the Sierra Nevada. It grows well at considerable distance from the ocean in New Zealand, France, Spain, and elsewhere (26,27).

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Damaging Agents

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Fire is the principal damaging agent in both young-growth and old-growth stands. The above-ground portions of young stands may be killed outright by a single ground fire, but the stands sprout and reoccupy the site. Fires are especially damaging to trees less than 20 years old because their thin bark does not protect them. Also, more flammable litter lies on the ground, and the microclimate is drier than under old-growth forest.

Old-growth redwood stands show evidence of three or more severe fires each century (23,44). In many instances, fires may only reduce the thickness of the protective bark, which may be more than 30 cm (12 in) thick. In other instances, fires cause basal wounds through which heart rots enter. The combination of recurring fires and advancing decay produces large basal cavities called "goose pens." In extreme instances, mature trees may be so weakened mechanically that they fall.

In its northern range, in and around Redwood National Park, CA, fire has a moderate ecological role in redwood stands. Light ground fires that do not open the canopy favor western hemlock regeneration but usually eliminate older hemlock from the stand. Douglas-fir establishment is infrequent and unsuccessful under a full overstory canopy, even following light ground fires on mesic sites. Relatively hot fires appear essential for the establishment of Douglas-fir trees in discrete age classes. Redwood, grand fir, and tanoak maintain their status in redwood stands with and without the influence of fire (47,48).

Frequency distributions of fires indicate a natural pattern of several short intervals between fires followed by one or more long interval. This suggests that prescribed burning to maintain ecosystems should also be done on a short-short-long interval pattern (23).

Redwood has no tree-killing diseases other than seedling diseases previously listed, but heart rots cause extensive cull. Most common heart rot in the southern part of the range of redwood is a brown cubical rot, caused by Poria sequoiae. Most significant farther north is a white ring rot caused by P. albipellucida (5,22).

A twig branch canker (Coryneum spp.) has been observed on sprouts and plantation trees of seedling and sapling size. This canker, which girdles stems and branches, could become damaging in plantations (5,22).

Several insects are found on redwood but none cause significant damage. These include a flatheaded twig borer and girdler (Anthaxia aeneogaster), two redwood bark beetles (Phloeosinus sequoiae and P. cristatus), and the sequoia pitch moth (Vespamima sequoiae) (21).

Bark stripping by the American black bear has caused serious damage in some parts of the redwood region. Wide strips of bark are ripped from the tree, often from the top to the ground, during April to August. Trees 10 to 30 years old and 15 to 25 cm (6 to 10 in) in diameter are damaged most and many may be girdled. Woodrats often injure planted trees on cutover land and occasionally attack sprouts and larger trees.

In a few instances, redwood is deformed by fasciation, a flattening of the normally cylindrical stem by formation of a row of linked meristems. The causes of most fasciations are unknown (40).

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Flowering and Fruiting

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Redwood is monoecious; inconspicuous male and female flowers are borne separately on different branches of the same tree. The ovulate conelets grow into broadly oblong cones (10). Redwood female strobili become receptive and pollen sheds between late November and early March, although flowering usually is over by the end of January. Weather conditions during pollination may directly affect seed quality. Continuous rains during flowering wash pollen from the male strobili and little pollen may reach the receptive female strobili. Dry periods during pollination permit better pollen dispersal and improve seed viability.

Redwood cones are terminal and are 13 to 29 mm (0.5 to 1.1 in) long. They mature in autumn of the first year after flowering and are open from early September until late December. Although cones persist for several months, they open and shed seeds soon after ripening.

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Genetics

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Sequoia is unique within Coniferales, being of a hexaploid nature (41). It was thought that redwood originated as an allopolyploid from hybrids between early Tertiary or Mesozoic species of Metasequoia and some extinct Taxodiaceous plant such as the modern giant sequoia. However, the types and numbers of marker chromosomes found in Metasequoia and Taxodium distichum are different than those present in Sequoia, making it unlikely that these species contributed to the polyploidy of Sequoia. Comparisons between the marker chromosomes in Sequoia and those in Sequoiadendron indicate that genomic contribution by Sequoiadendron to Sequoia is not probable (41).

Races of redwood are not known, but the following cultivars (cultivated varieties) have been recognized (16):

cv. 'Adpressa'  Tips of shoots creamy white. Awl-like leaves.
cv. 'Glauca'  Leaves 6 mm (0.25 in) long, glaucous, bluish.
cv. 'Nana Pendula'  Leaves glaucous, branches pendulous.
cv. 'Pendula'  Branches pendulous.
cv. 'Prostrata'  Prostrate at first; leaves green, glaucous beneath.

Four varieties of redwood now available in nurseries show a range of growth habits, texture, color, and form. They are named Aptos Blue, Los Altos, Soquel, and Santa Cruz (6).

An uncommon form of redwood, the albino redwood, has been described in a few locations within the redwood region (17). These albinos result from a genetic disorder and exist by attachment to a normal green tree, generally at the roots. The tallest albino observed was 19.8 m (65 ft) tall. Albinism is often a useful trait in genetics research to determine mutation rate, and for other purposes.

Preliminary results from studies of self and related outcross families indicate that, compared with outcrosses, selfing produced no additional cone abortion or variable effects on germination. Under stress conditions in nurseries and outplantings, some inbreeding depression becomes evident, and restricting inbreeding in redwood seed-orchards seems prudent (30).

The tissue culture techniques described earlier also allow genetic manipulation of redwood at the cellular level. Possibilities being explored include the production of di-haploid redwood from female gametophyte cultures (2).

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Growth and Yield

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Redwood is long lived, grows taller than any other tree species in the world, and is exceeded in bulk only by the giant sequoia. Redwoods are sexually mature at 10 years or less but continue to increase in volume for centuries. The oldest redwood found so far, determined by growth ring counts, is nearly 2,200 years old. Old-growth redwood forests sometimes are incorrectly called even-aged and overmature when, in fact, few forests in the world can match many redwood stands in range of ages and mixture of vigorously growing and decadent trees.

Redwood probably is best known for its great size, although the average redwood is smaller than commonly believed. Trees larger than 30 cm (12 in) in d.b.h. on a 12-ha (30-acre) old-growth tract in Humboldt County, CA, fell approximately into these divisions: 30 to 77 cm (12 to 30 in) in d.b.h., 50 percent; 78 to 153 cm (31 to 60 in), 32 percent; 155 cm (61 in) and larger, 18 percent. Redwoods 366 to 488 cm. (144 to 192 in) in d.b.h., found scattered over the entire range, are considered large. Trees 610 cm (240 in) or more in diameter at a point 1.5 m (5 ft) above the ground are rare.

Redwoods more than 61 m (200 ft) tall are common, and many trees growing on riverside benches, where soils are deep and moist, are taller than 91 m (300 ft). The tallest measured redwood was 112.1 m (367.8 ft) in 1964 (50).

Large trees and dense stocking combine to produce high yields. More than 81 percent of the commercial redwood forest land is classified as highly productive, and only 2 percent is poor for growing trees. Flats along rivers have yielded approximately 10,500 to 14,000 m³/ha (about 750,000 to 1,000,000 fbm/acre) in scaled logs. Harvest cuttings in Del Norte County, CA, on units of 5.3 ha (13 acres) and larger, produced gross volumes ranging from 1330 to 3921 m³/ha (95,000 to 280,000 fbm/acre, Scribner).

Biomass accumulates to record levels. A redwood stand in Humboldt State Park in California provides the greatest biomass ever recorded, with a stem biomass of 3461 t/ha (1,544 tons/acre) (20).

Economical conversion of old-growth redwood to young managed stands by shelterwood or selection cutting is difficult because net growth is negative during the decade after logging. Windthrow, slow growth of residual trees, and damage to established reproduction when residual trees are removed contribute to economic losses. Considering effect on growth, small clearcuttings seem to be a good method for converting old-growth redwood to young managed stands (9).

Young-growth redwood is often nearly as spectacular in size and yield as old growth. Dominant young-growth trees on good sites are 30.5 to 45.7 m (100 to 150 ft) tall at 50 years, and 50.3 to 67.1 m (165 to 220 ft) at 100 years. Height growth is most rapid up to the 35th year. On the best sites, however, height growth continues to be rapid well past 100 years (24,33).

Diameter growth of individual young trees can be rapid or extremely slow. In dense stands where competition is severe, annual diameter increment is commonly less than 1 mm (0.03 in). Occasionally, 40 or more rings per centimeter (more than 100/in) can be counted. At the other extreme, diameter growth sometimes exceeds 2.5 cm (1 in) a year. One redwood growing with little competition was 213 cm (84 in) in d.b.h. when 108 years old.

The yield of young-growth redwood stands at 100 years is expected to range from 742 m³/ha (10,600 ft³/acre) on low sites to 3576 m³/ha (51,080 ft³/acre) on high sites (32). The same stands yield 781 to 4998 m³/ha (55,760 to 357,000 fbm/acre International quarter-inch rule), and yields of more than 2800 m³/ha (about 200,000 fbm/acre International quarter-inch rule) are common in young-growth redwood stands. At earlier ages, however, the greatest yields are in stands that contain a mixture of redwood and Douglas-fir (25).

Natural pruning in young redwood stands often is not good. Although live crowns may be limited to the upper third of the trunk, dead limbs are persistent. Branch stubs, although decayed, may remain more than 50 years. In old trees, some branch stubs have affected the quality of the timber over a 200-year period. Trees in the intermediate crown class, however, often prune well naturally, and some trees in a heavily stocked stand have clean trunks for 23 to 30 m (75 to 100 ft) at 85 years.

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Reaction to Competition

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The redwood forest is a climax type. When growing with other species, redwood usually is a dominant tree. Douglas-fir can keep pace with redwood on many sites and occupy dominant and codominant crown positions along with redwood. Redwood has been classed as tolerant or very tolerant, the two highest categories in a scale of five shade tolerance classes. It is probably most accurately classed as very tolerant of shade in most situations.

Redwood stands are dense. At 60 years, redwood may have a basal area of more than 126 m²/ha (550 ft²/acre) on the best sites (32). Heavy stocking is desirable because the relatively high tolerance permits land to support a large number of dominant and codominant trees per unit area.

Under some conditions, redwood can endure suppression almost indefinitely. A 25-cm (10-in) suppressed tree might be more than 100 years old. Small trees may be suppressed for more than 400 years but still maintain a remarkable capacity to accelerate growth rates when released if they have not been crowded too closely and are not injured seriously during logging or slash burning. Large trees also can accelerate growth when released from competition.

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Rooting Habit

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Redwoods have no taproots, but lateral roots are large and wide-spreading. Small trees have better-than-average windfirmness, and large redwoods are windfirm under most conditions.

A study in extreme northwestern California indicated that a combination of wet soil and strong winds is necessary for significant windfall damage. Consequently, windfall is caused by only a few of the many winter storms. Storms that cause windfall come mainly from the south. Uprooting accounted for 80 percent of the redwood windfall in this study (7).

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Seed Production and Dissemination

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Redwoods start to bear seeds when 5 to 15 years old (8). One study showed that seed viability increased with the age of parent trees (38,39). Maximum seed viability was reached when trees were more than 250 years old. Seeds produced by trees younger than 20 years generally were less than 1 percent viable, and seeds from trees more than 1,200 years old were not more than 3 percent viable. Redwoods produce abundant seeds almost every year. Even trees in the intermediate crown class often produce seed crops. Fair to abundant crops were produced in 5 consecutive years in north coastal California (8). Cones often are rare, however, or nonexistent on large areas for many years in stands in Mendocino County, CA (central part of the range). Large mature stands on Maui, HI, have few or no cones or pollen (27).

Trees with new, narrow crowns resulting from sprouting of dormant buds after fire has killed the crown produce few cones during the first 4 years after the fire. About one-half such narrow-crowned trees, locally called fire-columns, bear cones in the fifth year, and almost all produce cones by the seventh or eighth year.

The germination rate of redwood seeds is usually low. Poor germination often results from a low percentage of sound seeds (less than 15 percent) rather than from dormancy. When obviously defective seeds are removed, germination rarely is below 80 percent, and is sometimes 100 percent (27). Identification of defective seeds often is difficult, however, because many seeds appearing sound are filled with tannin. In one seed study, soundness varied significantly with seed size. Seeds passing 12, 10, and 8 mesh screens were 2, 8, and 15 percent sound, respectively. Seeds from seven populations were photographed by X-ray. The distribution in categories was as follows: seeds empty or tannin filled, 58 to 97 percent; seeds from embryos damaged by fungi, 0 to 11 percent; and sound seeds, 1 to 32 percent (38,39).

Although only scant evidence is recorded on storage of redwood seeds, they do not seem to store well. One seed lot was stored successfully for 3 years but lost its viability completely after 5 years (19).

Redwood cones dry readily under conditions of low humidity and quickly release their seeds with slight shaking. But because weather conditions at cone ripening in nature usually are unfavorable for rapid drying, seed dispersal may be spread over periods that vary considerably in length. Rains, however, may hasten seed dissemination. One observer found in many instances that redwood seeds remained in the open cones until a drenching rain dissolved the tannic crystals in the cones (38,39). Seed dissemination during the winter months seems characteristic of redwood in the northern stands. More than four-fifths of the sound seeds in one study were shed during December and January.

Redwood seeds are small and light, number about 265,000/kg (120,000/lb), but lack efficient wings to slow them in falling (10). They fall at rates between 1.5 and 6.2 m/s (4.9 and 20.5 ft/s), averaging 2.6 m/s (8.6 ft/s). These rates are faster than for most other wind-disseminated forest seeds and limit seed dispersal considerably.

Timbered edges of clearcut units have effective seeding distances of only 61 m (200 ft) uphill and 122 m (400 ft) downhill under average redwood stand conditions. A recent study in Del Norte County, CA, showed that the largest clearcut units should not be more than 12 to 16 ha (30 to 40 acres) if regeneration will be completed by natural seeding (38,39). No silvicultural reasons exist for restricting the size of clearcuts, if areas are regenerated by artificial methods. Maximum size of clearcuttings is specified in Forest Practice Rules, based on erosion hazard, or other criteria.

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Seedling Development

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Redwood seeds, generally, are ready to germinate soon after they fall to the ground if seedbeds are moist and the weather is warm enough. Redwood seeds do not require pretreatment to germinate, but germination speed is increased by an overnight soak in aerated water (27). Mineral soil is the best seedbed, but seeds will germinate readily in duff, on logs, in debris, or under other vegetation, and in either shade or full sunlight if adequate soil moisture is available. Redwood seed germination is epigeal.

New redwood seedlings require a greater supply of soil moisture for survival than that needed by seedlings of most associated trees (19). Late spring and early fall rains can be critical survival factors. Apparently, redwoods have no root hairs. Consequently, redwood roots do not seem to function efficiently in extracting soil moisture. This fact may limit natural distribution to sites where favorable water relations result from high rainfall, humid air, moist soil, or low summer temperatures, or from various combinations of these conditions. Redwood seedlings on fully exposed soil can withstand considerable surface heat if their roots have reached a permanent moisture supply. Otherwise, they die before soil surface temperatures reach 60° C (140° F). Redwood seedlings are extremely vulnerable to infection by damping-off and Botrytis fungi during their first year (22).

In its early stages, redwood grows rapidly in height. Seedlings often grow about 46 cm (18 in) in the first season and trees 4 to 10 years old sometimes grow 0.6 to 2.0 m (2 to 6.5 ft) in a year. In many instances, however, rapid height growth of trees that originate from seed does not commence until the trees are more than 10 years old.

Juvenile growth of redwood is best in full sunlight. Although redwood seedlings can endure heavy shade, growth there is slow. Photosynthetic capacity in redwood is remarkably high at low light intensities and keeps increasing as light intensity increases, much like more intolerant species. Redwood grew vigorously in much weaker light than 12 other tree species studied (38,39). For example, it increased its size 8.8 times in 10 percent of full sunlight in a 9-month period, more than twice the growth of any of the other species in the test. For appreciable growth, Engelmann spruce (Picea engelmannii) and Douglas-fir require twice as much light as redwood. Pine requires three to four times as much.

Radial growth of redwood in Mendocino County, CA, at points 6, 14, and 32 km (4, 9, and 20 mi) from the coast did not vary markedly in growth pattern. Radial growth began after mid-March, increased to a maximum in late May, and then declined at a fairly uniform rate to a minimum at the end of September. Radial growth was negligible from October 1 to March 15.

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Soils and Topography

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The parent rock material of the redwood region is largely massive marine sandstone formed in the Tertiary and Upper Mesozoic periods. Considerable shale and lesser amounts of Mesozoic limestones and Franciscan slates, cherts, limestones, and sandstones also are present, and schists are fairly common in some localities.

High-site soils for redwood consist of Xerochrepts, Haploxerults, and Haplohumults of the Hugo, Josephine, Melbourne, Empire, Sites, and Larabee series (orders Inceptisols and Ultisols) and associated alluvial soils. The high-site residual soils have been derived from either consolidated or soft sedimentary rocks. In the Coastal Forest Practice Act District of California, which encompasses the natural range of redwood, the Hugo soil series predominates. In current soil taxonomic terms, the Hugo series is a Typic Distrochrept of the order Inceptisols (45,46). It is a member of a loamy-skeletal, mixed, mesic family, typically pale brown, moderately acid, gravelly (sandy) clay loam A horizons, and pale brown, strongly acid gravelly (sandy) clay loam B horizons. Limits of redwood forests sometimes are determined by soil types. For example, redwood does not grow on soils having high amounts of magnesium and sodium.

Fertility of soils under redwood stands has been studied by measuring the replaceable calcium concentration, expressed in equivalents, present in a square meter (10.76 ft²) to a depth of 30 cm (12 in). This measure indicates fertility best because it separates nutritional properties from other environmental effects. Equivalents ranged from 4 to more than 80, with 63 appearing to be optimum (49).

Soil nutrient levels that were observed to change during harvest of old-growth or second-growth redwood recovered to nearly original values during regrowth. In the one-meter soil profile, carbon, nitrogen, phosphorus, and exchangeable potassium and sodium increased in amount, while calcium decreased (52). Soil organic matter showed a small decline and recovery after logging (18).

The lowest amount of soil moisture available during the year has been related to minimum basal area growth of redwood stands. Basal area is used as an index of stand development. This minimum available soil moisture, expressed as a percentage of storage capacity, ranged between 18 and 86, with 62 correlated with maximum basal area (49).

The redwood region, generally, is characterized by irregular ridges oriented northwest to southeast with deep narrow valleys. Consequently, the principal streams drain to the northwest. Much of the terrain is rough, steep, and extremely dissected both by major streams and smaller drainages. Redwoods grow from sea level to about 915 m (3,000 ft) elevation, but most are found between 30 and 760 m (100 and 2,500 ft). The best stands have developed on flats and benches along the larger streams, on moist coastal plains, river deltas, moderate westerly slopes, and valleys opening toward the sea.

Although most redwood stands are close to the ocean, redwood does not tolerate ocean winds, and considerable evidence suggests that it is sensitive to ocean salts carried inland during storms. Usually redwoods do not grow on hillsides that face the ocean. The absence of redwood near the ocean also may be caused by the absence of forest soils of sufficient depth and fertility to support redwood.

Redwoods are smaller and give way to other species as altitude, dryness, and slope increase. In the north, redwoods clothe all exposures and reach their maximum development as forest trees. In the southern part of the range, redwoods are restricted to western or northern exposures, and at the extreme southern extension they are restricted almost entirely to the bottoms of narrow canyons that cut through steep foothills abutting the ocean. Trees near the mouths of these canyons often are exposed to onshore winds and frequently have flat tops with dead limbs on the windward side. This effect has been attributed to the trees' inability to replace moisture lost through desiccation by winds.

On alluvial flats, where redwoods reach their maximum development, soils have been built up by deposits of sediment from successive floods. In one area the ground level has been raised 3.4 m (11 ft) in 700 years. In another, repeated flooding in the past 1,000 years deposited nearly 9.1 m (30 ft) of silt and gravel around the bases of many large redwood trees. Deposits from a single flood have been as deep as 76 cm (30 in). Redwoods adapt to the new ground levels by originating new and higher root systems (43,51). This flooding generally kills competing species and thereby allows redwood to maintain nearly pure stands on such plains.

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Special Uses

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Redwood is used where decay resistance is important. Clark and Scheffer (14) found that decay resistance varied among trees or within the heartwood of individual trees. Decay resistance decreased from outer to inner hardwood. Wood classified as very decay resistant was about five times more prevalent in old-growth than in young-growth trees.

A prominent special feature of the redwood is its production of burls from which beautifully figured table tops, veneers, bowls, and other turned products are cut. These burls are found on any part of the trunk and in sizes varying from an inch to many feet in diameter. Their cause is unknown. Small burls containing hundreds of dormant buds often are cut and placed in shallow containers, kept moist, and allowed to sprout. These live burls serve as attractive house plants.

Another feature of redwood is its extremely tough and fibrous bark. The bark must be removed before logs reach the head saws so that sawing uniform lumber will be possible. The bark is used as hog fuel, insulation, or garden mulch.

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Vegetative Reproduction

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Redwood can be propagated by cuttings, but few large-scale attempts of this kind have been reported. In an early study in California, 40 percent of the cuttings from the tops of fast-growing seedlings that had been pushed into forest nursery soil with no special treatment developed root systems (38,39). Currently, rooting in excess of 90 percent is obtained routinely, with mist in a favorable medium, using juvenile cuttings from seedlings (27). Cuttings from older trees are more difficult to root.

Studies in the past 10 years have improved the cutting procedure by hedging-a technique that seems to maintain the juvenility of the donor tree. A single seedling and its clonal descendants can produce about 1 million cuttings in 3 years by repeated hedging of seedlings and their descendants (29).

Modern methods of plant tissue culture also have propagated redwood successfully (3). Tissues from outstanding mature trees may be cultured in nutrient medium, becoming undifferentiated masses of cells or callus. In different nutrient media, fragments of the callus can be induced to differentiate into small plants. When these plants become large enough, juvenile cuttings can be taken from them (30). In France, scientists have found that shoots of redwood 10 to 20 mm (0.4 to 0.8 in) long are the best reactive material for producing explants, with fragments of the annual shoots being more reactive than the annual sprouts of 2-year-old shoots (13). Tissue cultured plantlets are generally twice the size of seedlings of the same age (2).

Redwood can sprout from stumps and root crowns anytime of the year. Numerous and vigorous sprouts originate from both dormant and adventitious buds within 2 to 3 weeks after logging. Sprouting capacity is related to variables associated with tree size or age. Stumps of small young trees sprout more readily than those of large old trees (35). Stumps often are circled by more than 100 sprouts. Many sprouts may be necessary to sustain a healthy stump-root system (4,15). Powers and Wiant (37) found that sprout vigor was related to sprout density. Sprout vigor was reduced at densities less than one sprout per 2 feet of stump circumference. Each sprout soon develops its own root system, and in a remarkably short time the dominant sprouts create circles of new trees around the old stumps.

Depending on the intensity of thinning or partial cutting in redwood, sprouts grow and develop successfully in openings (11,31). A recent study showed that more than 90 percent of all redwood stumps sprouted in a 40-year-old redwood stand thinned to 25, 50, and 75 percent of the initial basal area. Consequently, all thinned stands contained several hundred redwood sprout clumps per acre, and several thousand individual sprouts. The heavier the thinning, the more sprouts developed into vigorous young crop trees (31).

Sprouting by redwood is principally from root crowns, but sprouts sometimes grow from the sides and tops of stumps. These high sprouts are less desirable because they are mechanically weak and not as vigorous as root-crown sprouts. Sprouts originating from the sides and top of stumps often are destroyed by strong wind.

Sprouts are commonly about 60 to 90 cm (24 to 36 in) tall at the end of the first year but may be more than 1.8 m (6 ft) tall. In one instance, a fire killed all sprouts around a stump. About 300 new sprouts appeared within a few days, and at the end of one growing season many reached 2.1 m (7 ft). Sprouts grow more rapidly than seedlings and the initial impetus lasts many years. However, the best phenotypes at age 40 to 80 in stands originating from both sprouts and seedlings often are found to be of seedling origin (27).

Early estimates of stocking from root crown sprouts varied from 20 to 35 percent of full stocking. A later study showed that redwood sprouts on old growth cutover redwood land in Mendocino and Humboldt Counties, CA, provided only 8 percent of full stocking. This finding is low compared to more recent stand examinations where the majority of redwood stems in 163 moderately to fully stocked young growth stands originated from sprouts (33).

Redwood can also sprout along almost the entire length of its trunk. If the crown of a tree is destroyed by fire or mechanically damaged, or the stem is suddenly exposed to light, numerous dormant buds along the trunk are stimulated and produce new foliage. Most of the trunk is then covered by feathery foliage extending 0.6 to 0.9 m (2 to 3 ft) from the trunk. Eventually, normal crowns develop again.

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Brief Summary

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Taxodiaceae -- Redwood family

David F. Olson, Jr., Douglass F. Roy, and Gerald A. Walters

Redwood (Sequoia sempervirens), also called coast redwood and California redwood, is native to the central and northern California coast, a region of moderate to heavy winter rain and summer fog so vital to this tree. It is one of three important North American trees of the family Taxodiaceae. Close relatives are the giant sequoia (Sequoiadendron giganteum) of the Sierra Nevada in California and the baldcypress (Taxodium distichum) of the southeastern states.

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Distribution

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The range of redwood extends southward from two groves on the Chetco River in the extreme southwest corner of Oregon (lat. 42° 09' N.), to Salmon Creek Canyon in the Santa Lucia Mountains of southern Monterey County, CA (lat. 35° 41' N.). This redwood belt is an irregular coastal strip about 724 km (450 mi) long and generally 8 to 56 km (5 to 35 mi) wide (39). Within this region, redwood trees grow now, or could grow, on an estimated 647 500 ha (1.6 million acres). Of this area, 260 200 ha (643,000 acres) comprise the commercial coast redwood forest type (more than 50 percent redwood stocking). The remainder of the area contains parks, other forest types containing redwood, and recently logged redwood type (12). The old-growth redwood, much of which is in State and National Parks, occupies less than 80 940 ha (200,000 acres) (36). The old-growth in commercial forests will be harvested within the next few decades. A major discontinuity splits the type in southern Humboldt County, CA. South of Sonoma County, CA, redwoods grow in detached and irregular areas to the southern extremity of the range (38,39).


- The native range of redwood.

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Physical Description

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Tree, Very large tree more than 75 m tall, Evergreen, Monoecious, Habit erect, Trees without or rarely having knees, Tree with bark rough or scaly, Young shoots in flat sprays, Buds not resinous, Leaves needle-like, Leaves scale-like, Leaves of two kinds, Leaves alternate, Needle-like leaf margins entire (use magnification), Leaf apex acute, Leaves < 5 cm long, Leaves < 10 cm long, Leaves not blue-green, Leaves white-striped, Scale leaves without raised glands, Needle-like leaves flat, Needle-like leaves not twisted, Needle-like leaf habit erect, Needle-like leaf habit drooping, Needle-like leaves per fascicle mostly 1, Needle-like leaf sheath early deciduous, Needle-like leaf sheath persistent, Twigs glabrous, Twigs not viscid, Twigs without peg-like projections or large fascicles after needles fall, Berry-like cones orange, Woody seed cones < 5 cm long, Bracts of seed cone included, Seeds tan, Seeds brown, Seeds winged, Seeds equally winged, Seed wings narrower than body.
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Sequoia sempervirens

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Trunk in sectional view
Sequoia sempervirensMHNT

Sequoia sempervirens (/səˈkwɔɪ.ə ˌsɛmpərˈvrənz/)[3] is the sole living species of the genus Sequoia in the cypress family Cupressaceae (formerly treated in Taxodiaceae). Common names include coast redwood, coastal redwood,[4] and California redwood.[5] It is an evergreen, long-lived, monoecious tree living 1,200–2,200 years or more.[6] This species includes the tallest living trees on Earth, reaching up to 115.9 m (380.1 ft) in height (without the roots) and up to 8.9 m (29 ft) in diameter at breast height. These trees are also among the longest-living organisms on Earth. Before commercial logging and clearing began by the 1850s, this massive tree occurred naturally in an estimated 810,000 ha (2,000,000 acres)[7][8][9] along much of coastal California (excluding southern California where rainfall is not sufficient) and the southwestern corner of coastal Oregon within the United States.

The name sequoia sometimes refers to the subfamily Sequoioideae, which includes S. sempervirens along with Sequoiadendron (giant sequoia) and Metasequoia (dawn redwood). Here, the term redwood on its own refers to the species covered in this article but not to the other two species.

Description

The coast redwood is known to have reached 115.5 m (379 ft) tall, with a trunk diameter of 9 m (30 ft).[10] It has a conical crown, with horizontal to slightly drooping branches. The trunk is remarkably straight. The bark can be very thick, up to 30 cm (1 ft), and quite soft and fibrous, with a bright red-brown color when freshly exposed (hence the name redwood), weathering darker. The root system is composed of shallow, wide-spreading lateral roots.

The leaves are variable, being 15–25 mm (58–1 in) long and flat on young trees and shaded lower branches in older trees. The leaves are scalelike, 5–10 mm (1438 in) long on shoots in full sun in the upper crown of older trees, with a full range of transition between the two extremes. They are dark green above and have two blue-white stomatal bands below. Leaf arrangement is spiral, but the larger shade leaves are twisted at the base to lie in a flat plane for maximum light capture.

The species is monoecious, with pollen and seed cones on the same plant. The seed cones are ovoid, 15–32 mm (9161+14 in) long, with 15–25 spirally arranged scales; pollination is in late winter with maturation about 8–9 months after. Each cone scale bears three to seven seeds, each seed 3–4 mm (18316 in) long and 0.5 mm (132 in) broad, with two wings 1 mm (116 in) wide. The seeds are released when the cone scales dry and open at maturity. The pollen cones are ovular and 4–6 mm (31614 in) long.

Its genetic makeup is unusual among conifers, being a hexaploid (6n) and possibly allopolyploid (AAAABB).[11] Both the mitochondrial and chloroplast genomes of the redwood are paternally inherited.[12]

Taxonomy

Scottish botanist David Don described the redwood as the evergreen taxodium (Taxodium sempervirens) in his colleague Aylmer Bourke Lambert's 1824 work A description of the genus Pinus.[13] Austrian botanist Stephan Endlicher erected the genus Sequoia in his 1847 work Synopsis coniferarum, giving the redwood its current binomial name of Sequoia sempervirens.[14] Endlicher probably derived the name Sequoia from the Cherokee name of George Gist, usually spelled Sequoyah, who developed the still-used Cherokee syllabary.[15] The redwood is one of three living species, each in its own genus, in the subfamily Sequoioideae. Molecular studies have shown that the three are each other's closest relatives, generally with the redwood and giant sequoia (Sequoiadendron giganteum) as each other's closest relatives.

However, Yang and colleagues in 2010 queried the polyploid state of the redwood and speculate that it may have arisen as an ancient hybrid between ancestors of the giant sequoia and dawn redwood (Metasequoia). Using two different single copy nuclear genes, LFY and NLY, to generate phylogenetic trees, they found that Sequoia was clustered with Metasequoia in the tree generated using the LFY gene, but with Sequoiadendron in the tree generated with the NLY gene. Further analysis strongly supported the hypothesis that Sequoia was the result of a hybridization event involving Metasequoia and Sequoiadendron. Thus, Yang and colleagues hypothesize that the inconsistent relationships among Metasequoia, Sequoia, and Sequoiadendron could be a sign of reticulate evolution (in which two species hybridize and give rise to a third) among the three genera. However, the long evolutionary history of the three genera (the earliest fossil remains being from the Jurassic) make resolving the specifics of when and how Sequoia originated once and for all a difficult matter—especially since it in part depends on an incomplete fossil record.[16]

Distribution and habitat

Coast redwoods occupy a narrow strip of land approximately 750 km (470 mi) in length and 8–75 km (5–47 mi) in width along the Pacific coast of North America; the most southerly grove is in Monterey County, California, and the most northerly groves are in extreme southwestern Oregon. The prevailing elevation range is 30–750 m (100–2,460 ft) above sea level, occasionally down to 0 and up to about 900 m (3,000 ft).[17] They usually grow in the mountains where precipitation from the incoming moisture off the ocean is greater. The tallest and oldest trees are found in deep valleys and gullies, where year-round streams can flow, and fog drip is regular. The terrain also made it harder for loggers to get to the trees and to get them out after felling. The trees above the fog layer, above about 700 m (2,300 ft), are shorter and smaller due to the drier, windier, and colder conditions. In addition, Douglas-fir, pine, and tanoak often crowd out redwoods at these elevations. Few redwoods grow close to the ocean, due to intense salt spray, sand, and wind. Coalescence of coastal fog accounts for a considerable part of the trees' water needs.[18] Fog in the 21st century is, however, reduced from what it was in the prior century, which is a problem that may be compounded by climate change.[19]

The northern boundary of its range is marked by two groves on mountain slopes along the north side of the Chetco River,[20] which is on the western fringe of the Klamath Mountains, near the California–Oregon border.[21][22] The northernmost grove is located within Alfred A. Loeb State Park and Siskiyou National Forest at the approximate coordinates 42°07'36"N 124°12'17"W. The southern boundary of its range is the Los Padres National Forest's Silver Peak Wilderness in the Santa Lucia Mountains of the Big Sur area of Monterey County, California. The southernmost grove is in the Southern Redwood Botanical Area, just north of the national forest's Salmon Creek trailhead and near the San Luis Obispo County line.[23][24]

The largest and tallest populations are in California's Redwood National and State Parks (Del Norte and Humboldt counties) and Humboldt Redwoods State Park, with the overall majority located in the large Humboldt County.

The prehistoric fossil range of the genus is considerably greater, with a subcosmopolitan distribution including Europe and Asia until about 5 million years ago. During the last ice age, perhaps as recently as 10,000 years ago, redwood trees grew as far south as the Los Angeles area (coast redwood bark found in subway excavations and at the La Brea Tar Pits).

Ecology

Fog is of major importance in coast redwood ecology. Redwood National Park.

The native area provides a unique environment with heavy seasonal rains up to 2,500 mm (100 in) annually. Cool coastal air and fog drip keep the forest consistently damp year round. Several factors, including the heavy rainfall, create a soil with fewer nutrients than the trees need, causing them to depend heavily on the entire biotic community of the forest, and making efficient recycling of dead trees especially important. This forest community includes coast Douglas-fir, Pacific madrone, tanoak, western hemlock, and other trees, along with a wide variety of ferns, mosses, mushrooms, and redwood sorrel. Redwood forests provide habitat for a variety of amphibians, birds, mammals, and reptiles. Old-growth redwood stands provide habitat for the federally threatened spotted owl and the California-endangered marbled murrelet.

Coast redwoods are resistant to insect attack, fungal infection, and rot. These properties are conferred by concentrations of terpenoids and tannic acid in redwood leaves, roots, bark, and wood.[25] Despite these chemical defenses, redwoods are still subject to insect infestations; none, however, are capable of killing a healthy tree.[25] Redwoods also face herbivory from mammals: black bears are reported to consume the inner bark of small redwoods, and black-tailed deer are known to eat redwood sprouts.[25]

The oldest known coast redwood is about 2,200 years old;[26] many others in the wild exceed 600 years. The numerous claims of older redwoods are incorrect.[26] Because of their seemingly timeless lifespans, coast redwoods were deemed the "everlasting redwood" at the turn of the century; in Latin, sempervirens means "ever green" or "everlasting". Redwoods must endure various environmental disturbances to attain such great ages.

In response to forest fires, the trees have developed various adaptations. The thick, fibrous bark of coast redwoods is extremely fire-resistant; it grows to at least a foot thick and protects mature trees from fire damage.[27][28] In addition, the redwoods contain little flammable pitch or resin.[28] If damaged by fire, a redwood readily sprouts new branches or even an entirely new crown,[27][25] and if the parent tree is killed, new buds sprout from its base.[25] Fires, moreover, appear to actually benefit redwoods by causing substantial mortality in competing species while having only minor effects on redwood. Burned areas are favorable to the successful germination of redwood seeds.[27] A study published in 2010, the first to compare post-wildfire survival and regeneration of redwood and associated species, concluded fires of all severity increase the relative abundance of redwood and higher-severity fires provide the greatest benefit.[29]

Redwoods often grow in flood-prone areas. Sediment deposits can form impermeable barriers that suffocate tree roots, and unstable soil in flooded areas often causes trees to lean to one side, increasing the risk of the wind toppling them. Immediately after a flood, redwoods grow their existing roots upwards into recently deposited sediment layers.[30] A second root system then develops from adventitious buds on the newly buried trunk and the old root system dies.[30] To counter lean, redwoods increase wood production on the vulnerable side, creating a supporting buttress.[30] These adaptations create forests of almost exclusively redwood trees in flood-prone regions.[25][30]

The height of S. sempervirens is closely tied to fog availability; taller trees become less frequent as fog becomes less frequent.[31] As S. sempervirens' height increases, transporting water via water potential to the leaves becomes increasingly difficult due to gravity.[32][33][34] Despite the high rainfall that the region receives (up to 100 cm), the leaves in the upper canopy are perpetually stressed for water.[35][36] This water stress is exacerbated by long droughts in the summer.[37] Water stress is believed to cause the morphological changes in the leaves, stimulating reduced leaf length and increased leaf succulence.[33][38] To supplement their water needs, redwoods use frequent summer fog events. Fog water is absorbed through multiple pathways. Leaves directly take in fog from the surrounding air through the epidermal tissue, bypassing the xylem.[39][40] Coast redwoods also absorb water directly through their bark.[41] The uptake of water through leaves and bark repairs and reduces the severity of xylem embolisms,[42][41] which occur when cavitations form in the xylem preventing the transport of water and nutrients.[41] Fog may also collect on redwood leaves, drip to the forest floor, and be absorbed by the tree's roots. This fog drip may form 30% of the total water used by a tree in a year.[37]

Reproduction

Redwood cone scales begin to open mid November, with seeds dispersing by the wind.
Basal regrowth after logging, Freshwater Creek (east of Eureka, California). While standing in the creek, a portion of the old stump can be seen.
View upward from the center of an old, logged stump (Freshwater Creek).

Coast redwood reproduces both sexually by seed and asexually by sprouting of buds, layering, or lignotubers. Seed production begins at 10–15 years of age. Cones develop in the winter and mature by fall. In the early stages, the cones look like flowers, and are commonly called "flowers" by professional foresters, although this is not strictly correct. Coast redwoods produce many cones, with redwoods in new forests producing thousands per year.[27] The cones themselves hold 90–150 seeds, but viability of seed is low, typically well below 15% with one estimate of average rates being 3 to 10 percent.[43][27] The low viability may discourage seed predators, which do not want to waste time sorting chaff (empty seeds) from edible seeds. Successful germination often requires a fire or flood, reducing competition for seedlings. The winged seeds are small and light, weighing 3.3–5.0 mg (200–300 seeds/g; 5,600–8,500/ounce). The wings are not effective for wide dispersal, and seeds are dispersed by wind an average of only 60–120 m (200–390 ft) from the parent tree. Seedlings are susceptible to fungal infection and predation by banana slugs, brush rabbits, and nematodes.[25] Most seedlings do not survive their first three years.[27] However, those that become established grow rapidly, with young trees known to reach 20 m (66 ft) tall in 20 years.

Coast redwoods can also reproduce asexually by layering or sprouting from the root crown, stump, or even fallen branches; if a tree falls over, it generates a row of new trees along the trunk, so many trees naturally grow in a straight line. Sprouts originate from dormant or adventitious buds at or under the surface of the bark. The dormant sprouts are stimulated when the main adult stem gets damaged or starts to die. Many sprouts spontaneously erupt and develop around the circumference of the tree trunk. Within a short period after sprouting, each sprout develops its own root system, with the dominant sprouts forming a ring of trees around the parent root crown or stump. This ring of trees is called a "fairy ring". Sprouts can achieve heights of 2.3 m (7 ft 7 in) in a single growing season.

Redwoods may also reproduce using burls. A burl is a woody lignotuber that commonly appears on a redwood tree below the soil line, though usually within 3 m (10 ft) in depth from the soil surface. Coast redwoods develop burls as seedlings from the axils of their cotyledon, a trait that is extremely rare in conifers.[27] When provoked by damage, dormant buds in the burls sprout new shoots and roots. Burls are also capable of sprouting into new trees when detached from the parent tree, though exactly how this happens is yet to be studied. Shoot clones commonly sprout from burls and are often turned into decorative hedges when found in suburbia.

Cultivation and uses

An example of a bonsai redwood, from the Pacific Bonsai Museum

Coast redwood is one of the most valuable timber species in the lumbering industry. In California, 3,640 km2 (899,000 acres) of redwood forest are logged, virtually all of it second growth.[1] Though many entities have existed in the cutting and management of redwoods, perhaps none has had a more storied role than the Pacific Lumber Company (1863–2008) of Humboldt County, California, where it owned and managed over 810 km2 (200,000 acres) of forests, primarily redwood. Coast redwood lumber is highly valued for its beauty, light weight, and resistance to decay. Its lack of resin makes it absorb water[44] and resist fire.

P.H. Shaughnessy, Chief Engineer of the San Francisco Fire Department wrote:

In the recent great fire of San Francisco, that began April 18th, 1906, we succeeded in finally stopping it in nearly all directions where the unburned buildings were almost entirely of frame construction, and if the exterior finish of these buildings had not been of redwood lumber, I am satisfied that the area of the burned district would have been greatly extended.

Because of its impressive resistance to decay, redwood was extensively used for railroad ties and trestles throughout California. Many of the old ties have been recycled for use in gardens as borders, steps, house beams, etc. Redwood burls are used in the production of table tops, veneers, and turned goods.

The Skyline-to-the-Sea Trail passing through a cut-out in a fallen California redwood tree

The Yurok people, who occupied the region before European settlement, regularly burned ground cover in redwood forests to bolster tanoak populations from which they harvested acorns, to maintain forest openings, and to boost populations of useful plant species such as those for medicine or basketmaking.[25]

Extensive logging of redwoods began in the early nineteenth century. The trees were felled by ax and saw onto beds of tree limbs and shrubs to cushion their fall.[25] Stripped of their bark, the logs were transported to mills or waterways by oxen or horse.[25] Loggers then burned the accumulated tree limbs, shrubs, and bark. The repeated fires favored secondary forests of primarily redwoods as redwood seedlings sprout readily in burned areas.[25][45] The introduction of steam engines let crews drag logs through long skid trails to nearby railroads,[25] furthering the reach of loggers beyond the land nearby rivers previously used to transport trees.[45] This method of harvesting, however, disturbed large amounts of soil, producing secondary-growth forests of species other than redwood such as Douglas-fir, grand fir, and western hemlock.[25] After World War II, trucks and tractors gradually replaced steam engines, giving rise to two harvesting approaches: clearcutting and selection harvesting. Clearcutting involved felling all the trees in a particular area. It was encouraged by tax laws that exempted all standing timber from taxation if 70% of trees in the area were harvested.[25] Selection logging, by contrast, called for the removal 25% to 50% of mature trees in the hopes that the remaining trees would allow for future growth and reseeding.[45] This method, however, encouraged growth of other tree species, converting redwood forests into mixed forests of redwood, grand fir, Sitka spruce, and western hemlock.[45][25] Moreover, the trees left standing were often felled by windthrow; that is, they were often blown over by the wind.

The coast redwood is naturalized in New Zealand, notably at Whakarewarewa Forest, Rotorua.[46] Redwood has been grown in New Zealand plantations for more than 100 years, and those planted in New Zealand have higher growth rates than those in California, mainly because of even rainfall distribution through the year.[47]

Other areas of successful cultivation outside of the native range include Great Britain, Italy, Portugal,[48] Haida Gwaii, middle elevations of Hawaii, Hogsback in South Africa, the Knysna Afromontane forests in the Western Cape, Grootvadersbosch Forest Reserve near Swellendam, South Africa and the Tokai Arboretum on the slopes of Table Mountain above Cape Town, a small area in central Mexico (Jilotepec), and the southeastern United States from eastern Texas to Maryland. It also does well in the Pacific Northwest (Oregon, Washington, and British Columbia), far north of its northernmost native range in southwestern Oregon. Coast redwood trees were used in a display at Rockefeller Center and then given to Longhouse Reserve in East Hampton, Long Island, New York, and these have now been living there for over twenty years and have survived at 2 °F (−17 °C).[49]

This fast-growing tree can be grown as an ornamental specimen in those large parks and gardens that can accommodate its massive size. It has gained the Royal Horticultural Society's Award of Garden Merit.[50][51]

Statistics

Dried resin

Fairly solid evidence indicates that coast redwoods were the world's largest trees before logging, with numerous historical specimens reportedly over 122 m (400 ft).[52]: 16, 42  The theoretical maximum potential height of coast redwoods is thought to be limited to between 122 and 130 m (400 and 427 ft), as evapotranspiration is insufficient to transport water to leaves beyond this range.[32] Further studies have indicated that this maximum requires fog, which is prevalent in these trees' natural environment.[53]

A tree reportedly 114.3 m (375 ft) in length was felled in Sonoma County by the Murphy Brothers saw mill in the 1870s,[54] another claimed to be 115.8 m (380 ft) and 7.9 m (26 ft) in diameter was cut down near Eureka in 1914,[55][56] and the Lindsey Creek tree was documented to have a height of 120 m (390 ft) when it was uprooted and felled by a storm in 1905. A tree reportedly 129.2 m (424 ft) tall was felled in November 1886 by the Elk River Mill and Lumber Co. in Humboldt County, yielding 79,736 marketable board feet from 21 cuts.[57][58][59] In 1893, a Redwood cut at the Eel River, near Scotia, reportedly measured 130.1 m (427 ft) in length, and 23.5 m (77 ft) in girth.[60][61][62] However, limited evidence corroborates these historical measurements.

Today, trees over 60 m (200 ft) are common, and many are over 90 m (300 ft). The current tallest tree is the Hyperion tree, measuring 115.61 m (379.3 ft).[26] The tree was discovered in Redwood National Park during mid-2006 by Chris Atkins and Michael Taylor, and is thought to be the world's tallest living organism. The previous record holder was the Stratosphere Giant in Humboldt Redwoods State Park at 112.84 m (370.2 ft) (as measured in 2004). Until it fell in March 1991, the "Dyerville Giant" was the record holder. It, too, stood in Humboldt Redwoods State Park and was 113.4 m (372 ft) high and estimated to be 1,600 years old. This fallen giant has been preserved in the park.

As of 2016, no living specimen of other tree species exceeds 100 m (330 ft).

The largest known living coast redwood is Grogan's Fault, discovered in 2014 by Chris Atkins and Mario Vaden in Redwood National Park,[26] with a main trunk volume of at least 1,084.5 cubic meters (38,299 cu ft)[26] Other high-volume coast redwoods include Iluvatar, with a main trunk volume of 1,033 m3 (36,470 cu ft),[52]: 160  and the Lost Monarch, with a main trunk volume of 988.7 m3 (34,914 cu ft).[63]

Albino redwoods are mutants that cannot manufacture chlorophyll. About 230 examples (including growths and sprouts) are known to exist,[64] reaching heights of up to 20 m (66 ft).[65] These trees survive like parasites, obtaining food from green parent trees. While similar mutations occur sporadically in other conifers, no cases are known of such individuals surviving to maturity in any other conifer species. Recent research news reports that albino redwoods can store higher concentrations of toxic metals, going so far as comparing them to organs or "waste dumps".[66][67]

List of tallest trees

Heights of the tallest coast redwoods are measured yearly by experts.[26] Even with recent discoveries of tall coast redwoods above 100 m (330 ft), it is likely that no taller trees will be discovered.[26]

Diameter is measured at 1.4 m (4 ft 7 in) above average ground level (at breast height). Details of the precise locations for most of the tallest trees were not announced to the general public for fear of causing damage to the trees and the surrounding habitat.[26] The tallest coast redwood easily accessible to the public is the National Geographic Tree, immediately trailside in the Tall Trees Grove of Redwood National Park.[68]

List of largest trees

The following list shows the largest S. sempervirens by volume known as of 2001.[52]: 186–7 

Calculating the volume of a standing tree is the practical equivalent of calculating the volume of an irregular cone,[69] and is subject to error for various reasons. This is partly due to technical difficulties in measurement, and variations in the shape of trees and their trunks. Measurements of trunk circumference are taken at only a few predetermined heights up the trunk, and assume that the trunk is circular in cross-section, and that taper between measurement points is even. Also, only the volume of the trunk (including the restored volume of basal fire scars) is taken into account, and not the volume of wood in the branches or roots.[69] The volume measurements also do not take cavities into account. Most coast redwoods with volumes greater than 850 m3 (30,000 cu ft) represent ancient fusions of two or more separate trees, which makes determining whether a coast redwood has a single stem or multiple stems difficult.[70] Starting in 2014, more record-breaking coast redwood trees were discovered. The largest disclosed was a massive redwood called Grogan's Fault/Spartan,[71] which has been measured to have a volume of 38,300 cubic feet.

Details of the precise locations for most of the tallest trees were not announced to the general public for fear of causing damage to the trees and the surrounding habitat.[26] The largest coast redwood easily accessible to the public is Iluvatar, which stands prominently about 5 meters (16 ft) to the southeast of the Foothill Trail of Prairie Creek Redwoods State Park.

Other notable examples

See also

References

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Sequoia sempervirens: Brief Summary

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Trunk in sectional view Sequoia sempervirens – MHNT

Sequoia sempervirens (/səˈkwɔɪ.ə ˌsɛmpərˈvaɪrənz/) is the sole living species of the genus Sequoia in the cypress family Cupressaceae (formerly treated in Taxodiaceae). Common names include coast redwood, coastal redwood, and California redwood. It is an evergreen, long-lived, monoecious tree living 1,200–2,200 years or more. This species includes the tallest living trees on Earth, reaching up to 115.9 m (380.1 ft) in height (without the roots) and up to 8.9 m (29 ft) in diameter at breast height. These trees are also among the longest-living organisms on Earth. Before commercial logging and clearing began by the 1850s, this massive tree occurred naturally in an estimated 810,000 ha (2,000,000 acres) along much of coastal California (excluding southern California where rainfall is not sufficient) and the southwestern corner of coastal Oregon within the United States.

The name sequoia sometimes refers to the subfamily Sequoioideae, which includes S. sempervirens along with Sequoiadendron (giant sequoia) and Metasequoia (dawn redwood). Here, the term redwood on its own refers to the species covered in this article but not to the other two species.

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