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裸子植物
Pinus ponderosa Douglas ex P. Lawson & C. Lawson
EOL Text
Cope's giant salamander is found in the Puget lowland forests along with several other western North America ecoregions. The Puget lowland forests occupy a north-south topographic depression between the Olympic Peninsula and western slopes of the Cascade Mountains, extending from north of the Canadian border to the lower Columbia River along the Oregon border. The portion of this forest ecoregion within British Columbia includes the Fraser Valley lowlands, the coastal lowlands locally known as the Sunshine Coast and several of the Gulf Islands. This ecoregion is within the Nearctic Realm and classified as part of the Temperate Coniferous Forests biome.
The Puget lowland forests have a Mediterranean-like climate, with warm, dry summers, and mild wet winters. The mean annual temperature is 9°C, the mean summer temperature is 15°C, and the mean winter temperature is 3.5°C. Annual precipitation averages 800 to 900 millimeters (mm) but may be as great as 1530 mm. Only a small percentage of this precipitation falls as snow. However, annual rainfall on the San Juan Islands can be as low as 460 mm, due to rain-shadow effects caused by the Olympic Mountains. This local rain shadow effect results in some of the driest sites encountered in the region. Varied topography on these hilly islands results in a diverse assemblage of plant communities arranged along orographically defiined moisture gradients. Open grasslands with widely scattered trees dominate the exposed southern aspects of the islands, while moister dense forests occur on northern sheltered slopes characterized by Western red cedar (Thuja plicata), Grand fir (Abies grandis), and Sword fern (Polystichum munitum) communities.
There are only a small number of amphibian taxa in the Puget lowland forests, namely: Cope's giant salamander (Dicamptodon copei); Monterey ensatina (Ensatina eschscholtzii); Long-toed salamander (Ambystoma macrodactylum); Western redback salamander (Plethodon vehiculum); Northwestern salamander (Ambystoma gracile); Pacific chorus frog (Pseudacris regilla); Coastal giant salamander (Dicamptodon tenebrosus); Rough-skin newt (Taricha granulosa); the Vulnerable Spotted frog (Rana pretiosa); Tailed frog (Ascopus truei); and Northern red-legged frog (Rana aurora).
Likewise there are a small number of reptilian taxa within the ecoregion: Common garter snake (Thamnophis sirtalis); Western terrestrial garter snake (Thamnophis sirtalis); Northern alligator lizard (Elgaria coerulea); Western fence lizard (Sceloporus occidentalis); Northwestern garter snake (Thamnophis ordinoides); Sharp-tailed snake (Contia tenuis); Yellow-bellied racer (Coluber constrictor); and Western pond turtle (Clemmys marmorata).
There are numberous mammalian taxa present in the Puget lowland forests. A small sample of these are:Creeping vole (Microtus oregoni), Raccoon (Procyon lotor), Southern sea otter (Enhydra lutris), Mink (Mustela vison), Coyote (Canis latrans), Black-tailed deer (Odocoileus hemionus), Pallid bat (Antrozous pallidus), and Harbour seal (Phoca vitulina).
A rich assortment of bird species present in this ecoregion, including the Near Threatened Spotted owl (Strix occidentalis), Turkey vulture (Cathartes aura), Bald eagle (Haliaeetus leucocephalus), Blue grouse (Dendragapus obscurus), as well as a gamut of seabirds, numerous shorebirds and waterfowl.
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Rights holder/Author | cc-by-nc-sa 3.0 |
Source | http://www.eoearth.org/view/article/51cbeebf7896bb431f699fd6/54dfd31a0cf21994e423f4cf/?topic=51cbfc79f702fc2ba8129ee0 |
Trees to 72m; trunk to 2.5m diam., straight; crown broadly conic to rounded. Bark yellow- to red-brown, deeply irregularly furrowed, cross-checked into broadly rectangular, scaly plates. Branches descending to spreading-ascending; twigs stout (to 2cm thick), orange-brown, aging darker orange-brown, rough. Buds ovoid, to 2cm, fully 1cm broad, red-brown, very resinous; scale margins white-fringed. Leaves 2--5 per fascicle, spreading to erect, persisting (2--)4--6(--7) years, 7--25(--30)cm ´ (1--)1.2--2mm, slightly twisted, tufted at twig tips, pliant, deep yellow-green, all surfaces with evident stomatal lines, margins serrulate, apex abruptly to narrowly acute or acuminate; sheath 1.5--3cm, base persistent. Pollen cones ellipsoid-cylindric, 1.5--3.5cm, yellow or red. Seed cones maturing in 2 years, shedding seeds soon thereafter, leaving rosettes of scales on branchlets, solitary or rarely in pairs, spreading to reflexed, symmetric to slightly asymmetric, conic-ovoid before opening, broadly ovoid when open, 5--15cm, mostly reddish brown, sessile to nearly sessile, scales in steep spirals (as compared to Pinus jeffreyi ) of 5--7 per row as viewed from side, those of cones just prior to and after cone fall spreading and reflexed, thus well separate from adjacent scales; apophyses dull to lustrous, thickened and variously raised and transversely keeled; umbo central, usually pyramidal to truncated, rarely depressed, merely acute, or with a very short apiculus, or with a stout-based spur or prickle. Seeds ellipsoid-obovoid; body (3--)4--9mm, brown to yellow-brown, often mottled darker; wing 15--25mm.
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Rights holder/Author | eFloras.org Copyright © Missouri Botanical Garden |
Source | http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200005351 |
Tree, Evergreen, Monoecious, Habit erect, Trees without or rarely having knees, Tree with bark rough or scaly, Young shoots 3-dimensional, Buds resinous, Leaves needle-like, Leaves alternate, Needle-like leaf margins finely serrulate (use magnification or slide your finger along the leaf), Leaf apex acute, Leaves > 5 cm long, Leaves < 10 cm long, Leaves > 10 cm long, Leaves yellow-green above, Leaves yellow-green below, Leaves not blue-green, Needle-like leaves somewhat rounded, Needle-like leaves twisted, Needle-like leaf habit erect, Needle-like leaves per fascicle mostly 2, Needle-like leaves per fascicle mostly 3, Needle-like leaf sheath persistent, Twigs glabrous, Twigs viscid, Twigs not viscid, Twigs without peg-like projections or large fascicles after needles fall, Berry-like cones orange, Woody seed cones > 5 cm long, Seed cones bearing a scarlike umbo, Umbo with obvious prickle, Bracts of seed cone included, Seeds brown, Seeds yellow, Seeds winged, Seeds unequally winged, Seed wings prominent, Seed wings equal to or broader than body.
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Rights holder/Author | Compiled from several sources by Stephen C. Meyers, Oregon State University in collaboration with Aaron Liston, Oregon State University, Steffi Ickert-Bond, University of Alaska Fairbanks, and Damon Little, New York Botanical Garden. |
Source | http://plants.usda.gov/java/profile?symbol=PIPO |
Ponderosa pine is monoecious. At pollination the male strobili, borne in short, dense clusters, are 2 to 3 cm (0.8 to 1.2 in) long and female conelets are 2.5 cm (1 in) long. In western Montana, central Idaho, and eastern Oregon, at elevations from 910 to 1830 m (3,000 to 6,000 ft), flowering generally begins between May 1 and 10. Pollen is shed May 25 to June 15, cones reach a full size of 8 to 15 cm (3 to 6 in) July 20 to August 10 of the next year, seed is ripe August 20 to September 5, cones begin to open September 1 to 13, and seed is shed until November. On the east and west sides of the Sierra Nevada in California, at an elevation of 1830 m (6,000 ft), however, cones develop about 2 weeks later (13). In northern Arizona, near Flagstaff, pollen is shed between June 10 and 20 (55), but at an elevation of 910 m (3,000 ft) on the west slope of California's Sierra Nevada, pollen has been collected as early as April 15; May 11 was average for a 7-year period. Also on the west slope of the Sierra Nevada, pollen is shed an average of 8 days later for each 300 m (1,000 ft) rise in elevation (13).
In Colorado, at 2710-m (8,900-ft) elevation, during a 9-year period, female conelets emerged on or about June 18 and only about 36 percent of them survived until the beginning of the second year. Flowering is correlated closely with the passing of freezing weather (13).
Ponderosa pine is propagated by seed. Cones are ready for collection in October and November when they turn reddish brown. Mature seed is firm and brown in color. Cones should be dried on canvas tarp in a well-ventilated area immediately after they have been collected. The seeds will drop from the cones as they dry.
Several germination methods for ponderosa pine have been utilized, each with their own variations. In general, seeds undergo an imbibation treatment before stratification. Seeds are placed in mesh bags and soaked in cold running water for 48 hours. One variation is to soak the seeds in a 40% bleach solution for 10 minutes with hand agitation prior to placing them under running water. The mesh bags are place in plastic bags and stored at 1oC for 2 to 8 weeks. They should be checked daily for mold. Seeds are sown into containers and covered with media. The media should be kept moist throughout germination. Germination will occur at an average greenhouse temperature of 20oC. Alternating greenhouse temperatures of 21-25oC during the day and 16-18oC at night is an appropriate environment for germinating seeds. Germination will occur in approximately 15 days.
Seedlings are thinned and watered daily throughout the establishment phase. They should not be moved outdoors until after the last frost of the year.
Seeds can be dried to between 5 and 8% moisture and placed in airtight plastic bags, then stored for long periods of time in freezers set at –15oC.
Trees to 70 m tall; trunk to 2.5 m d.b.h. in native range; bark yellow- to red-brown, deeply and irregularly furrowed into broadly oblong, scaly plates; crown broadly conical to rounded; branchlets orange-brown, aging darker, stout, rough; winter buds red-brown, ovoid, very resinous, scales white fringed at margin. Needles tufted at apex of branchlets, spreading to erect, (2 or)3(-5) per bundle, deep yellow-green, slightly twisted, 7-25(-30) cm × (1-)1.2-2 mm, pliant, stomatal lines present on all surfaces, base with persistent sheath 1.5-3 cm, margin serrulate. Seed cones solitary or rarely paired, sessile or subsessile, mostly reddish brown, broadly ovoid when open, symmetric or asymmetric, 5-15 cm, maturing in 2 years, then soon shedding seeds, leaving rosettes of scales on branchlets. Apophyses dull or lustrous, thickened, variously raised, cross keeled; umbo usually pyramidal or truncate, rarely depressed or with a reflexed prickle. Seeds brown or yellow-brown, often mottled darker, ellipsoid-obovoid, 3-9 mm; wing 1.5-2.5 cm.
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Rights holder/Author | eFloras.org Copyright © Missouri Botanical Garden |
Source | http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=200005351 |
Ponderosa pine grows to impressive size. Stems 263 cm (103.5 in) in d.b.h. and 70.7 m (232 ft) in height have been recorded (13). Diameters at breast height of 76 to 127 cm (30 to 50 in) and heights of 27.4 to 39.6 m (90 to 130 ft) are common throughout most of its range. Trees often reach ages of 300 to 600 years.
Diameter growth can be rapid and remain fairly constant for long periods provided trees are given adequate growing space. In California, on productive sites, free-growing trees can reach 66 cm (26 in) in d.b.h. in 30 years or 22 cm. (8.7 in) per decade (data on file at Pacific Southwest Forest and Range Experiment Station, Redding, CA). In central Oregon, where sites are less productive, trees 13 to 51 cm (5 to 20 in) in d.b.h. and from 19 to 36 years old can grow 12 cm (4.9 in) in d.b.h. per decade if free of intertree competition (3). Trees in a virgin stand in Arizona grew 29 mm (1.14 in) on the average during a 10-year period, but trees in a cutover stand grew 43 mm (1.68 in) (55).
Vegetative competition can restrict diameter growth markedly whether it be from neighboring trees or understory shrubs. In the central Oregon study, trees completely surrounded by understory shrubs grew only 9 cm (3.5 in) per decade. Those trees with no competitive ground cover averaged 12 cm (4.7 in) of growth per decade. In California on a droughty, skeletal soil, severe shrub competition reduced diameter growth to less than half that of competition-free trees. Insect damage, which was greater on the trees competing with shrubs, accounted for some of the growth depression (44). Stagnation in diameter, and often in height, represents a serious problem in densely stocked stands throughout the species' range, but especially on poor sites.
Height growth is most rapid in the pole and young sawtimber size classes to about 60 years. In the Pacific Northwest, dominant trees in stands of moderate density grow from 0.24 to 0.46 m (0.8 to 1.5 ft) annually between the ages of 20 to 60 years on timber-producing sites (2). Rate of growth declines gradually at older ages. Arizona trees of 160 years or older (determined at breast height) grow little in height (55). Height growth increases with site productivity and is more sensitive to stand density than was once believed.
Representative yields of ponderosa pine from a normal yield table for sites of various productivities are given in table 1 (39). For extensive natural stands, table values should be reduced by 25 percent or more because of roads, rock outcrops, steep slopes, openings, and other unproductive areas.
Table 1- Total volume inside bark of ponderosa pine 1.5 cm (0.6 in) and larger in d.b.h. (39) Site index at base age 100 years¹ Age 18 m or 60 ft 27 m or 90 ft 37 m or 120 ft 46 m or 150 ft yr m³/ha 20 28 94 168 262 40 122 238 396 588 60 192 340 570 861 80 238 413 696 1060 100 273 472 794 1204 120 308 518 868 - 140 336 556 928 - yr ft³/acre 20 400 1,350 2,400 3,750 40 1,750 3,400 5,650 8,400 60 2,750 4,850 8,150 12,300 80 3,400 5,900 9,950 15,150 100 3,900 6,750 11,350 17,200 120 4,400 7,400 12,400 - 140 4,800 7,950 13,250 - ¹Height of dominant and codominant trees of average d.b.h. Old-growth ponderosa pine produces clear, high-grade lumber, but young trees typically are limby. Natural pruning develops slowly. An average clear length of only 3.5 m (11.5 ft) was recorded in 250-year-old stands in central Idaho (13).
Ponderosa pine can be over-irrigated in poorly drained soils, or drowned out on high water table sites.
It responds well to thinning, which should be done as stands become older to develop larger crowns, resulting in heavier seed crops for wildlife. More forage for deer and elk become available from associated plants by opening the canopy. The use of repellents or other control measures may be necessary to prevent overuse of the trees by rodents.
Ponderosa pine is resistant to fire due to its thick bark. Low intensity surface fires control competitive species like scrub oak and shade-tolerant conifers. Ponderosa pine seedlings can also survive low intensity burns.
Habitat and Ecology
Systems
- Terrestrial
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Rights holder/Author | International Union for Conservation of Nature and Natural Resources |
Source | http://www.iucnredlist.org/apps/redlist/details/42401 |
Population Differences Ponderosa pine shows distinct geographic variations over its widespread range. Within and between var. ponderosa and var. scopulorum, provenance studies (51,65,66) have shown genetic variation in growth, stem form, needle length, survival, initiation of leader growth, seasonal pattern of root growth potential, ability to germinate under moisture stress (41), biotic and abiotic damage (17,26,52), monoterpene production (58), nutrient status (29,68), and isozymes (10). This wealth of information on genetic diversity was summarized and interpreted recently (10). It suggests that var. ponderosa consists of three major geographic races and var. scopulorum of two major geographic races. Within var. ponderosa, the Pacific race is found in California northward from the Transverse Ranges and west of the Sierra Nevada and Cascade Range into northern Oregon. Pacific race pines have relatively large needles, cones, and seeds, and are rapid growing and least cold hardy in tests to date. The North Plateau race extends northward along the eastside of the Sierra Nevada and Cascade Range and east to the Continental Divide in Montana. Like the Pacific race, it has open, plume-like foliage, 3-needle fascicles and isozyme characteristics. But the North Plateau race has needles with thickened layers of hypoderm and sunken stomata, and is indistinguishable from the Rocky Mountain race in monoterpene characteristics. Least well understood, but distinct in monoterpene production, is the Southern California race.
Within var. scopulorum, the Rocky Mountain race comprises the northeast portion of the species' range. It is characterized by compact foliage, 2-needle fascicles, and better growth in trials east of its natural range. The Rocky Mountain race joins the Southwestern race along a broad, ill-defined front through southern Colorado, Utah, and Nevada. The Southwestern race has relatively open foliage, low proportions of 2-needle fascicles, and resins with distinctive monoterpene composition.
Results from a provenance study of 45-year-old trees in northern Idaho and a study of 30-year-old trees in Oregon and Washington (60) showed that 36 percent of the variation in the height of the trees was associated with seed source. A clinal variation was evident in the increase of height from sources in an east-to-west direction. This variation was related to September-through-June precipitation. Clinal variation in a latitudinal and altitudinal direction was related to April-May temperatures. Incidence of animal damage and of frost injury was related, also, to seed source.
Ponderosa pine varies markedly in its resistance to cold. In a test of 298 individual tree progenies planted in Michigan, all 2-year-old seedlings of California origin suffered severe injury from cold (66). Progenies from British Columbia, Washington, eastern Oregon, Arizona, and southern New Mexico suffered light damage. No damage was reported for progenies from the remainder of the species' range. Essentially the same pattern was found in the northern Idaho study in 10- to 15-year-old trees (65).
Elevational. variation has been studied intensively in central Idaho (53) and in California (9). On the west slope of the Sierra Nevada in California, seeds collected from trees growing at elevations of 40 to 2130 m (125 to 7,000 ft) were planted at altitudes of 290, 830, and 1720 m (950, 2,730, and 5,650 ft) above sea level. The general trend was that early growth was most rapid for mid-elevation sources and least rapid for high-elevation sources, regardless of the elevation of the plantation. But by 29 years, at the high-elevation plantation, sources from high elevations had overtaken sources from low elevations and had nearly caught up to sources from middle elevations. Middle and low elevation sources, especially the latter, suffered stem and leader damage from snow and wind, which significantly reduced their growth superiority. Wood specific gravity decreased with increasing elevation of parent source regardless of where the source was planted (16). No elevational effect was discerned in tracheid length, although individual differences were found. Differences were recognized, also, in total height and diameter, and in the seasonal growth pattern (42) for families within elevational zones. Genetic diversity among populations, both in California and central Idaho, was readily interpretable as adaptive variation. Results of both studies suggest that for selective breeding of a wide-ranging species with distinct elevational differentiation, such as ponderosa pine, superior progenies can be obtained from selection within the optimum elevational zone of best geographic sources. In central Idaho, the recommended elevational zone is ± 180 m (600 ft).
Hybrids Natural crosses of ponderosa pine with Jeffrey pine have been observed in California where their ranges overlap, but they are rare. Where the two species grow in the same stand, different flowering times and other reproductive barriers restrict crossing (11). Ponderosa pine crosses with Pinus montezumae and P. arizonica, and rarely with P. engelmannii (45). Introgressive hybridization has been observed with P. washoensis.
In addition to the natural hybrids, artificial crosses have been obtained with a number of other hard pine species, including P. durangensis.