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California Oak Woodland is a "conglomeration" of range types unified mostly by the annual species in the understory of these various types. Baker in Barbour and Major (1995, p. 384) outlined the California oak woodland range types. In an abbreviated or condensed format these are:
The Society for Range Management recognized and described only three of these seven oak woodland types including the riparian one. This left only the blue oak and coast live oak phases receiving SRM coverage.
1. California blue oak (Quercus douglasii)-annual grass woodland- This predominately woodland community with an annual grass understory could logically be lumped in with either the Savanna or Forest and Woodland slides depending on canopy cover, or interpretation of the individual vegetation scientist, forester, or rangeman. So too could those communites of California annual grassland with widely scattered blue oaks be seen as grassland, savanna, or woodland with any guidelines being at best subjective. In fact this very situation has lead to claims and counterclaims by the various public agencies (eg. California Division of Forestry, California Agricultural Extension Service) amounting to legal and philosophical challenges as to who has authority, responsibility, expertise, etc. for management of this vegetation depending on whether these communities are forests, grasslands, or whatever. Using some arbitrary percentage of oak canopy cover or quantity for tree stocking (growing space determined by criteria of density or occupancy) could lead to different jurisdictions by several organizations for specific oak stands or oak-grass communities side-by-side in the same pasture!
This landscape-scale scene of California blue oak-dominated range in the foothills of the central Sierra Nevada is from an altitude of 28,000-30,000 feet. It shows the variation in tree density and canopy coverage typical of this vegetation. It was included here in the Forest and Woodland slides based on tree dominance and physiogonomy, two major standards for type designation and mapping. The Society for Range Management (Shiflet, 1994) designated this range cover type as "Blue Oak Woodland (201)". For consistency and deference to the authority/expertise of the SRM these California blue oak-annual grass communities were included with Forest and Woodland slides. In this Sierra foothills landscape, areas of annual grassland of considerable acreage are clearly visible. Examples of California annual grassland range, including some that are as much woodland as grassland, were included with the Grassland slides.
Placer County, California. Estival aspect, June. California blue oak-dominated communities: FRES No. 28 (Western Hardwoods Forest Ecosystem), K-26 (California Oakwoods). SAF 250, but the "pure blue oak" phase or "blue oak savanna" described therein (Eyre, 1980, ps.126-127). Kuchler (1977) California vegetation map unit No. 26 (Blue Oak-Digger Pine Forest), but again pure blue oak form or phase. California annual grassland: FRES No. 42 (Annual Grasslands Ecosystem), but no K- number for annual grassland. SRM 201 for Blue Oak Woodland and SRM 215 for California annual grassland (SRM title of Valley Grassland, but most is Coast Range or Sierra foothills grassland as the valley was converted to agronomic and horticultural crops). Encinal (Oak) Series in California Evergreen Forest and Woodland biotic community of Brown et al. (1998).
2. Blue oak-annual grass woodland- Two consociations comprise this grass- woodland community. California blue oak is the only tree species present and it forms a woodland structure with crowns close but not touching each other. Wild oats (Avena fatua) grows in a "pure" (single-species) stand to form the grass understory except on shallow soils and dry south slopes where, as seen in the center of this slide, soft chess or soft brome (Bromus mollis) is about the only grass. Redstem filaree (Erodium cicutarium) is an associate of soft chess on the south slopes.Research and observation has conclusively shown that wild oats is a decreaser and soft brome is a decreaser or increaser depending on range site in the California annual range type. Based on species composition and quantity of mulch (weight of herbage remaining at end of grazing season) this annual grass range was managed perfectly. Yolo County, California. Estival aspect, June.
The photographs of annual grass-woodland range (both here and in the Grassland slides) show that the understory is essentially "grassland" with the oak crowns comprising interrupted overstories which distinguish the communities as woodlands or savannas distinct from grassland in a strict sense. The understory is the same in all cases (annual grass-dominated) with potential species composition varying by range site. FRES No. 28 (Western Hardwoods Ecosystem), K-26 (California Oakwoods) and the understory is FRES No. 42 (Annual Grasslands Ecosystem), with no Kuchler unit for California annual grassland (the Kuchler unit 41 [Californian Steppe] is for the "extinct" Pacific perennial bunchgrass prairie). Blue oak variant of SAF 250 (Blue Oak-Digger Pine) for overstory; SRM 201 (Blue Oak Woodland) but the understory is the same as SRM 215 (Valley Grassland) or California Native Plant Society California Annual Grassland Series. Encinal (Oak) Series in California Evergreen Forest and Woodland biotic community of Brown et al. (1998); would be the Quercus douglasii Association (if such was listed) to be consistent with the Brown et al. (1998) classification system..
3. California blue oak-annual grass woodland- This is a "picture-perfect" example of this range cover type in Excellent range condition class, and with a healthy multi-age population of blue oak. The annual grass understory is dominated by wild oats with ripgut brome (Bromus rigidus = B. diandrus) "second in command" (main associate). Research over the last two-thirds of a century by various workers has consistently shown tht thest two taller, larger annual grasses are the main two decreasers of the anthropogenic climax annual grass range. Ripgut requires attentive grazing management . Livestock are best taken off by time of seed set and development of the mechanically injurious callus of the lemma. The high yield of herbage and greater nutritive value of these two Eurasian annuals justifies management of the range for them as key species. Abundance of characteristic persistent glumes of wild oats and the amount of mulch remaining on these relatively steep slopes at end of the grazing/grass- growing season indicates excellent management of this obviously productive range.
The various age classes of blue oak are well represeted indicating at least some oak regeneration. As is the case with the beloved valley or California white oak (Quercus lobata), blue oak reproduction has been inadequate for regeneration (mortality exceeds natality) with severe declines in populations of these valuable trees. Major research and extension efforts are under way (eg. University of California Hardwood Task Force) to reverse this unsatisfactory trend. Even in this picturesque scene blue oak seedling establishment is nil.
Tehama County, California. Estival aspect, June. FRES No. 28 (Western Hardwoods Ecosystem), K-26 (California Oakwoods), California blue oak variant of SAF 250 (BlueOak-Digger Pine). SRM 201 (Blue Oak Woodland). California Native Plant Society Blue Oak Series. Quercus douglasii Association in Encinal (Oak) Series, California Evergreen Forest and Woodland biotic community of Brown et al. (1998).
4. Interior live oak (Quercus wislizenii)- annual grass woodland- This is the interior live oak-dominated phase of foothill oak woodland. It is not a single-species form like the California blue oak phase. The woodland in this photograph includes some blue oak, Pacific madrone (Arbutus mensiesii), and in deeper draws on north slopes California black oak. Tree stocking is overwhelmingly interior live oak. There is a shrub layer made up of large individuals of common manzanita (Arctostaphylos manzanita), California buckeye (Aesculus californica) and, on edges between tree canopy and open grassland, blue elderberry (Sambucus coerulea). The dominant shrub in openings is chamise and the dominant grass of grassland and woodland understory is wild oats (both of these latter species are distinguishible in right foreground). Other common herbs are soft chess or soft brome, redstem filaree, red brome (Bromus rubens), and silver hairgrass (Aira caryophyllea).
University of California Hopland Field Station, Mendocino County, California. Estival aspect, June. Oak woodland is: FRES No. 28 (Western Hardwoods Ecosystem), K-26 (California Oakwoods), interior live oak variant of SAF 250 (Blue Oak-Digger Pine), SRM 201 (Blue Oak Woodland), California Native Plant Society Interior Live Oak Series. California annual grassland is: FRES No. 42 (Annual Grasslands Ecosystem), no Kuchler designation, SRM 215 (Valley Grassland), California Native Plant Society California Annual Grassland Series. Mixed Quercus Association in Encinal (Oak) Series or, alternatively, a northern variant of the Mixed Hardwood Series in California Evergreen Forest and Woodland biotic community of Brown et al. (1998).
5. Mixed oak-annual grass woodland- Two views of a combination of blue oak and interior live oak phases of California oak woodland with a California annual grassland understorey. The oak-annual grass woodland is most precisely described and envisioned as a merging of open oak forest and naturalized annual grassland: a "hybrid" type in which annual grassland comprises the understorey of an oak-dominated canopy in which tree crowns do not interlock or contact each other except at rareintervals. In some instances trees are so widely scattered that physiogonomy appears more as a grassland with occasional trees (ie. a textbook example of savanna).
Oak-annual grass woodland sometimes has yet another mixture or "hybridized" component when there are several oak (and perhaps other tree) species with no single species being an obvious dominant. In other words, the oaks do not fall clearly into any of the recognized California Oak Woodland phases cited above. The vegetation shown in these two photographs was such an oak woodland community. Blue oaks were more numerous than other tree species, but the largest trees were interior live oak. In addition there were individual trees of California black oak (Quercus kelloggii) and Oregon white oak (Q. garryana). There was almost no regeneration of any of the oak species.
A shrub layer was lacking and there were but rare individuals (trace portions) of common manzanita, blue elderbery, coast red elderberry (Sambucus callicarpa) and, even more limited, California buckeye . The latter two woody species were closer to size and form of small trees so as to produce limited browse.
The herbaceous understorey was well developed and composed almost entirely of high successional status grass species. The dominant species was wild oats. Soft brome or soft chess was the herbaceous associate. The amber patches were populated largely by soft chess whereas the green areas consisted primarily of wild oats with some ripgut brome. Little quaking grass (Briza minor) grew in pure, dense, but very small patches. There were smaller proportions of other annual grasses of low successional status including fescue (Festuca dertonensis), mouse barley (Hordeum leporinum), red brome, dogtail grass (Cynosurus echinatus), and silver hairgrass. Redstem filaree was the most important forage-producing forb.
This particular range plant community was in a large exclosure that had been protected from livestock grazing for 30 years. As a result the grass component was dominated by the decreasers (wild oats and ripgut brome) or the sensitive increaser soft chess (grazing response of this species from decreaser to increaser depending on range sites).
Foster Biological Area, University of California Hopland Field Station Mendocino County, California. Late May. Oak woodland is: FRES No. 28 (Western Hardwood Ecosystem), K-26 (California Oakwoods),.mixed oak variant of SAF 250 (Blue Oak-Digger Pine), SRM 201 (Blue Oak Woodland), California Native Plant Society combination of Blue Oak and Interior Live Oak Series. California annual grassland is: FRES No. 42 (Annual Grasslands Ecosystem), no Kuchler designation, SRM 215 (Valley Grassland), California Native Plant Society Annual Grassland Series. Mixed Quercus Association in Encinal (oak) Series or, alternatively, a northern variant of the Mixed Hardwood Series in California Evergreen Forest and Wooodland biotic community of Brown et al. (1998).
6. Excellent herbaceous understorey of mixed oak-annual grass woodland- This range of this annual grass-blue oak-interior liveoak range had been grazed more or less continuously by sheep for decades. For five years up to time of this photograph this herbaceous understorey had been grazed at 100% of moderate calibrated stocking rates. Range condition class was Excellent. The dominant Eurasian and naturalized annual grass on this particular oak woodland range was wild oats, but on many local spots (microsite scale), especially depressions an north slopes, ripgut brome was dominant (as shown in this photograph of the understorey sward) Ripgut is probably the most most sensitive important decreaser on California annual range. Soft chess and redstem filaree, two other Eurasian species that produce high-quality range feed, were also common on this pasture. Dominance of wild oats and ripgut demonstrated proper grazing management (stocking rate in particular) and the potential for high forage production of annual grass-oak woodland range.
These dominant grass species were shown and discussed herein under California grasslands. Plant and animal responses during a five-year stocking rate trial on this and two other annual grass-oak woodland ranges in the California Coast Range were reported by Rosiere (1987) and Rosiere and Torell (1996).
Range S3, University of California Hopland Field Station. Mendocino County, California. Late May (peak standing crop). California annual grassland portion of annual grass-oak woodland is: FRES No. 42 (Annual Grasslands Eocsystem), SRM 215 (Valley Grassland), California Native Plant Society Annual Grassland Series.
7. California Blue Oak-Digger Pine Foothill Woodland- This is the classic foothill woodland of the Pacific side of the Sierra Nevada. It is the Pinus-Quercus Association of the Pine-Oak Woodland of Clements (1920, p. 202-204). Elevationally it occurs immediately above the single-species blue oak-dominated woodland. Where the blue oak phase of foothill woodland is absent the blue oak-Digger pine type is the first zone of woody vegettion above California annual grassland of the valley and foot slope. In the example seen here the herbaceous understory is a continuation of annual grassland species, often tending to grasses of smaller, more xeric species like soft brome, red brome, silver hairgrass, foxtail fescue (Festuca megalura= Vulpia myuros var. hirsuta) rather than the larger dominants of valley grassland like wild oats and ripgut brome. Forbs are more common as well (eg. the filarees). Shrubs are those of the lower slopes like common manzanita, whiteleaf manzanita (Arctostaphylos viscida), scrub interior live oak (Quercus wislizenii var, frutescens), blue elderberry, California buckeye (obvious in this shot), wedgeleaf ceanothus (Ceanothus cunneatus), toyon (Photinia arbutifolia), and, in draws and north slopes, coyotebrush (Baccharis pilularis), poison oak (Rhus diversiloba), and California coffeeberry (Rhamnus californica). There are a few interior live oak, but California black oak is absent. This example is a very diverse range community because trees are widely spaced and the woodland extends from a shallow stream below up to the beginning of the Sierran Lower Montane Forest with an interior draw that has pronounced east and west slopes. The blue-gray plant colony in left foreground is the horribly noxious weed, yellow star-thistle (Centaurea solstitialis).
Butte County, California. Estival aspect, June. FRES No. 28 (Western Hardwoods Ecosystem), K-26 (California Oakwoods), classic form of SAF 250 (Blue Oak-Digger Pine), SRM 201 (Blue Oak Woodland), combination of California Native Plant Society Blue Oak and Foothill Pine Series. Oak-Pine Series in California Evergreen Forest and Woodland biotic community of Brown et al. (1998).
8. Digger Pine, known also as gray pine, ghost, and foothill pine (Pinus sabiniana)- This is one of the western yellow pines (Harlow et al. 1979, p. 113-114). It was named after the Digger Indians for whom the seeds were second only to acorns as a plant food source (Peattie, 1953, p. 94). In contrast to many of the Indian tribes of eastern and central North America the aborigines of the Great Basin and California did not develop technology or master the Spanish horse to such extent as to "wax fat" from their natural resources. Instead their generally primitive lives were a day-to-day hard-scramble existence. Even Digger pine nuts seem less substantive than those of the pinyon pine of the Rocky Mountain Region. Digger was a derogatory term used generically for several tribes (eg. Piute) after their habit of digging in the ground for roots, rodents, etc. As one watches modern-day Californians face power shortages because they were too short-sighted and environmentally "green" to wisely plan for future energy needs he appreciates the dignity, resourcefulness, and capacity for life of the first Californians. This reminds the student of natural resources that man, no matter how technologically backward or advanced, is the manipulator of ecosystems:
"What is man, that thou art mindful of him? and the son of man, that thou visitest
him? For thou hast made him a little lower than the angels…Thou madest him to
have dominion over the works of thy hands; thou hast put all things under his feet…"
Psalms 8: 4-6 King James Version
Peattie (1953, p. 94) pointed out that Digger pine often grows at right angles to the predominant slope such as to lean away from the hillside and the pull of gravity giving it a picturesque outline. The cones which are the second largest, heaviest of the American pines remain attached to the tree and open slowly over a number of years. The Indians gathered the somewhat serotinous cones and placed them in fires forcing them to yield the pine nuts (Sudworth, 1967, p. 54). Note in this shot the prominent California wild grape (Vitis californica) climbing the pines. Jepson (1953) remarked that this could kill trees, but they typically co-exist. The California wild grape is readily eaten by deer (Sampson and Jespersen, 1963, p. 116). Poor ole Diggers could have a picnic here!
9. Mixed Evergreen Foothill Woodland- This is the Digger Pine-Canyon Live Oak (Quercus chrysolepis)-Interior Live Oak form of the general pine-oak woodland. Specifically, it is the elevational subtype immediately above the California blue oak phase and is one form of the interior live oat phase (Griffin in Barbour and Major, 1995, p. 401). The view here is of a southwestern slope. The understory is made up of species of the California annual range type, it being an extension or continuation of valley grassland. Butte County, California. Estival aspect, June. FRES No. 28 (Western Hardwoods Ecosystem), K-26 (California Oakwoods), variant of SAF 250 (Blue Oak- Digger Pine), variant of SRM 201 (Blue Oak Woodland), combination of California Native Plant Society Foothill Pine and Canyon Live Oak Series. Oak-Pine Series in California Evergreen Forest and Woodland biotic community of Brown et al. (1998).
California Mixed Evergreen Forests is a collective term that encompasses a set or array of Pacific coastal mountain forest communities. These are spatially situated within (ie. "sandwiched" between) the Northern Pacific Coast Coniferous Forests (the complex of Redwood Forest, Cedar-hemlock-Douglas Fir Forest, Spruce-Cedar-Hemlock Forest, etc.) to the west and California Oak-Pine Woodland, California Annual Grassland, and various forms of California Chaparral to the east. This general community of mixed evergreen forest forms often occurs as a mosaic within these types (Sawyer et al. in Barbour and Major, 1995, ps. 360-361). It extends from southern Oregon to southwestern California, but it's form as zonal vegetation is limited mostly to central and, especially, northern sections.
10. Diagram of zones of vegetation along the elevational gradient of the central Sierra Nevada- Source: Shelford (1963, p. 170). Communities along this generalized transect vary depending on latitude or even general section of the Sierra (eg. northern, central, southern Sierra Nevada) as well as factors like soils, slope, aspect, etc.
Oregon white oak or Garry oak (Quercus garryana) forms range plant communities in which this is the only tree species present or at least the only one of any meaningful contribution or cover except where unnatural fire exclusion allows invasion by such species as Douglas fir (Pseudotsuga menziesii). Density and cover of Oregon white oak in these one-tree species communities under natural conditions is more that of a woodland or savanna than of a forest form which develops under protection from fire (Franklin and Dryness, 1973, ps. 126-129). In the interior valleys of Oregon Oregon white oak woodland exist as different communities differentiated by which shrub species dominant the understorey (Franklin and Dryness, 1973, ps. 111-114). Successional patterns of Oregon white oak communities were noted in these parts of Franklin and Dryness (1973).
Among forest and range ecosystems recognized by the U.S. Forest Service (Garrison et al., 1977) the Western Hardwoods Ecosystem (FRES No. 28) consisted of the two Kuchler units of Oregon Oakwoods (K-22) and California Oakwoods (K-26). This distinction seemed obvious given dominance of Oregon oak communities by a single species and that of the California oak types by several species (and frequently more than one Quercus species in each type which carried the designation of phase as listed above).
11.Perimeter or edge of Oregon white oak woodland- The edge of a Garry or Oregon white woodland that adjoined a naturalized Coast Range grassland had mature trees with the horizonal spreading limbs and open crown characteristic of trees that developed in uncrowded growing spaces. This was actually closer to the savanna rather than the woodland and, ceertainly, the forest form of this species.
The understorey was herbaceous and mostly a single layer of primarily perennial (with some annual), introduced, cool-season grasses. In this photograph the Eurasian perennials, tall fescue (Festuca arundinacea) and orchardgrass (Dactylis glomerata) were dominant, but the native blue wildrye (Elymus galucus) was present as a relict species of the pre-Columbian vegetation.
Benton County, Oregon. June. Estival aspect. FRES No. 28 (Western Hardwoods Forest Ecosystem). K-22 (Oregon Oakwoods). SAF 233 (Oregon White Oak). Oregon White Oak Series in Oregonian Deciduous and Evergreen Forest biotic community or regional formation of Brown et al. (1998). Oregon white oak/blue wildrye association of Kagan et al. (2004). Willamette Valley-Valley Foothills Ecoregion, 3d (Pater, undated).
12. Savanna or open woodland of Oregon white oak and naturalized grasses in Oregon Coast Ranges- Physiognomy of a savanna-like form of the what Kuchler (1964, 1966) called the Oregon Oakwoods. The understorey was herbaceous being dominated by introduced and naturalized perennial and annual, cool-season, Eurasian grasses. Most of the perennial grass species were introduced first for agronomic pasture and later bred for turf selections as production of turfgrass seed became a major industry throughout the Willamette Valley. The annual grasses included several of the Mediterranean species that comprised the naturalized (disclimax) annual grasslands of the California Coast Range and Central Valley. Thus this savanna or woodland community was a combination of numerous species from several similar expressions of vegetation (inclding related range types).
There were scattered small trees (smaller than saplings) of invading Pacific Douglas-fir. This was genesis of a second tree layer that in absence of fire will probably eventually replace the Oregon White Oaks. These Douglas-fir were visible in the left background beneath scattered oaks. Succession in plant communities of the Coast Ranges and interior valleys of Oregon has been limited to relatively few studies. These were reviewed and general tentative conclusions published by Franklin and Dyrness (1973, ps. 126-129). The concensus appeared to be that numerous climax communities were possible depending on range or forest site.
Benton County, Oregon. June. Estival aspect. FRES No. 28 (Western Hardwoods Forest Ecosystem). K-22 (Oregon Oakwoods). SAF 233 (Oregon White Oak). Oregon White Oak Series in Oregonian Deciduous and Evergreen Forest biotic community of Brown et al. (1998). Oregon white oak/blue wildrye association of Kagan et al. (2004). Willamette Valley-Valley Foothills Ecoregion, 3d (Pater, undated).
13. Clump of Oregon white oaks in a savanna of Oregon white oak and naturalized Eurasian grasses- This assemblage of Oregon white oaks was growing along the edge of a white oak woodland adjacent to a naturalized grassland composed primarily of Eurasian perennial and annual grasses. It was on a ridge top within the interior Coast Ranges of northern Oregon. Presence of large numbers of horizonal limbs (= primary branches) very close to the trunk base indicated that these trees had grown and developed free of the shading influence of other trees. This branching pattern or form was more indicative of a savanna or, at least woodland, tree cover rather than that of a forest.
The understorey was exclusively herbaceously except for some regeneration of Oregon white oak (seedling or very young oak in center foreground). Perennial grasses included tall fescue, orchardgrass, and colonial bentgrass (Agrostis capillaris). Annual grasses present were dogtail grass or hedgehog dogtail (Cynosurus echinatus) and soft chess or soft brome (Bromus mollis). Native blue wildrye was locally common or even dominant, but overall it was an associate of the naturalized Eurasian perennial grasses that dominated the herbaceous layer.The most common forb was the large and, again, introduced Eurasian species, common burdock (Arctium minus).
Benton County, Oregon. June. Estival aspect. FRES No. 28 (Western Hardwoods Forest Ecosystem). K-22 (Oregon Oakwoods). SAF 233 (Oregon White Oak). Oregon White Oak Series in Oregonian Deciduous and Evergreen Forest biotic community or regional formation of Brown et al. (1998). Oregon white oak/blue wildrye association of Kagan et al. (2004). Willamette Valley-Valley Foothills Ecoregion, 3d (Pater, undated).
14. Interior of Oregon white oak woodland in Oregon Coast Ranges- These two views were inside an Oregon or Garry oak stand with tree stocking of such density as to constitute the forest (vs. woodland and savanna) form of this cover type. For all of its simple beauty this range plant community was botanically diverse with regard to both species richness and structure. Plants present varied from non-vascular epiphytes like lichen and mosses (note patches of moss on tree trunks) to numerous grass species to widely scattered shrubs (including two roses [Rosa species]) and forbs like common burdock.
Tree trunks that were free of lower limbs indicated the forest form of this forest range cover type in contrast to the numerous horizonal limbs of the savanna and woodland forms presented above. The herbaceous understorey consisted overwhelming of perennial grasses. At the edge of this Oregon white oak-dominated community there was a greater cover and density of annual grassses (though introduced and native perennial grass species were still predominant) than in the more shaded interior of this forest range community. The grassy understorey shown in the first slide was composed primarily of introduced agronomic species like tall fescue, orchardgrass, colonial bentgrass, and perennial ryegrass (Lolium perenne) with scattered bunches of native blue wildrye. Grass understorey in the second slide consisted of these same speciesexcept that blue wildrye was dominant. Vegetation of this second slide was interpreted as climax or pre-Columbian (ie. it was the pontential natural vegetation).
Regeneration of Oregon white oak was evident in both of these "photo-plots". Recruitment consisted of trees of varying sizes but sapling-size was most prevalent. There were also a few snags of mid-size oaks the cause of death of which was unknown.
Readers should appreciate the potential of the grazing resource of this Oregon white oak woodland. Incidentially, "woodland" or "oakwoods" has been used by workers in naming and describing this cover type or ecosystem without distinction as to forest and woodland in the strict sense. Strictly speaking woodland is most commonly applied to "a plant community in which, in contrast to a typical forest, the trees are often small, characteristically short-boled relative to their crown depth, and forming only an open canopy with the intervening area being occupied by lower vegetation, commonly grass" (Helms, 1998).Woodland has often been specified to be a tree-dominated plant community with trees so widely spaced that their crows do not interlock or form a continuous canopy where forest refers to tree-dominated vegetation that at climax and/or with mature trees develops a closed or continuous canopy. Both forests and woodlands in a strict snese are "woodlands" in generic usage.
The difference between forest and woodland may depend exclusively on the distinctions between crown size and shape or canopy cover (ie. spacing among individual trees or tree density) or there may be species differences (particularily in lower layers) in vegetation. The savanna expression or form of a tree-dominated (at least aspect dominance or physiognomy) community has trees that are even more widely spaced or having lower density and aerial cover. Plant species composition may be the same or different in comparisons among savanna, forest, and woodland, and savannar.
The same Oregon white oak community was featured in most of the photographs in this section. By strict definition savanna, woodland, and forest forms were all present. The savanna pattern was also present at landscape scale where relatively small stands of Garry oak were scattered across a large expanse of naturalized grassland. In context of Landscape Ecology the generic Oregon white oak woodland functioned as patches in the matrix of grassland.
Benton County, Oregon. June. Estival aspect. FRES No. 28 (Western Hardwoods Forest Ecosystem). K-22 (Oregon Oakwoods). SAF 233 (Oregon White Oak). Oregon White Oak Series in Oregonian Deciduous and Evergreen Forest biotic community of Brown et al. (1998). Oregon white oak/blue wildrye association of Kagan et al. (2004).
15. Interior of climax Oregon white oak woodland range- Two views of pristine Garry or Oregon white oak woodland in the Coast Ranges of northern Oregon. Blue wildrye was an almost exclusive dominant in the vegetation shown here as to be an understorey consociation. In contrast to understorey vegetation shown in some earlier photographs where introduced agronomic grasses were dominant, here blue wildrye (the native dominant decreaser grass) continued to rule supreme. The vegetation presented in these two slides and in the immediately preceding set of two slides was interpreted as the climax range plant community. It represented the virgin vegetation at its natural potential.
Viewers can readily detect the conspicuous spike inflorescences of blue wildrye.
Lack of a woody layer or even minor component (other than a few native roses) was obvious and an exception to many Garry oak woodland communities (Franklin and Dyrness, ps. 111-115). Invasion by occasional Pacific Douglas-fir (sapling-size) was mentioned above and discussed below.
Viewers should also note in the second slide the Oregon white oak snag (right-most trunk in foreground). Previous photographs showed regeneration of Oregon white oak. In this "snapshot" the on-going population dynamics, the life-and-death of individuals of a species, was captured at a point in time. Vegetation is never static; it is dynamic. This was the fundamental lesson taught to generations of rangemen, foresters, and other students of Vegetation Science by Henry Chandler Cowls and Frederic Edward Clements. This dynamic nature of vegetation funcitions at various spatial and temporal scales for both individual species and the whole plant community. Stay tuned for the next set of slides.
Benton County, Oregon. June. Estival aspect. FRES No. 28 (Western Hardwoods Forest Ecosystem). K-22 (Oregon Oakwoods). SAF 233 (Oregon White Oak). Oregon White Oak Series in Oregonian Deciduous and Evergreen Forest biotic community of Brown et al. (1998). Oregon white oak/blue wildrye association of Kagan et al. (2004). Willamette Valley-Valley Foothills Ecoregion, 3d (Pater, undated).
16. Invasion of Oregron white oak woodland or on-going succession? The dynamic nature of vegetation was mentioned in context of species population dynamics in the preceding set of slides. In this set, vegetation dynamics was portrayed at the plant community level of organization. Pacific Douglas-fir had begun invading the Oregon white oak-dominated range vegetation. This range plant community was interpreted as the climax or potential natural vegetation. So where does Douglas-fir enter in (no pun intended)? One of the more readily available --if dated-- sources that described plant succession in the Oregon white oak community has been Franklin and Dyrness (1973, ps. 126-129). These authors concurred with earlier investigators and concluded that there were several pathways of plant succession for this community ranging from climax (steady state) Oregon white oak forests to replacement of Garry oak by Pacific Douglas-fir and/or bigleaf maple. In historical retrospect, the discussioin by Franklin and Dyrness (1973, ps. 126-129) could be seen as a forerunner of the state and transition concept (models) of succession.
Description of the Oregon white oak forest cover type by the Society of American Foresters (Eyre, 1980, p. 110-111) was very similar to that of Franklin and Dyrness (1973) except that Eyre (1980, p. 110-111) considered the role of fire (both natural and aboriginal) in maintenance of the Oregon white oak type. The Society of American Foresters description stated that fire had been critical in maintaining open woodland and savanna forms of Garry oak-dominated communities.
Obviously periodic surface fires-- as almost certainly occurred in the pre-Columbian vegetation-- would prevent long-term establishment of Douglas-fir. The fire regimen (especially season and frequency, but perhaps also intensity) might differ where the grassy understorey was dominated by introduced and naturalized Eurasian grasses rather than native species like blue wildrye. On the other hand it is possible that there would be little difference because both exotic, agronomic perennials and native perennial bunchgrasses are cool-season and primarily festucoid grasses. An understorey dominated by Mediterranean annual grasses would logically alter fire regimen(s) more than would Eurasian perennial grasses.
Designation of Douglas-fir establishment as invasion (or as an invader or invading species) was used in the original Clementsian meaning: "Invasion is the movement of one or more plants from one area into another and their establishment in the latter. It is thus the complete and complex process of which migration, ecesis, and competition are the essential parts. It is going on at all times and in all directions" (Weaver and Clements, 1938, p. 166). Invasion can be either plant succession, the forward progression of vegetation development along the sere, or retrogression, the retrograde movement in vegetation back down the sere toward a lower stage of plant community development. Retrogression on range and forest often results in invasion of climax or late seral vegetation by nonclimax woody and herbaceous species that direct vegetation to a lower seral stage. This is known as range depletion. Under other management, conditions of resource use, and cover types or sites invasion by, say, woody plants (climax or nonclimax) can be on-going succession and further plant community development toward climax.
So is the invasion of the Oregon white oak woodland by Douglas-fir the natural progression of plant succession or is it retrogression? Is there an absolute answer to this question? If Oregon white oak woodland is but a seral stage to a Pacific Douglas-fir forest climatic climax (in the Clementsian model) the answer would seem to obvious. Upon further thought, however, this conclusion would lead to acceptance of fire as a part of climate. If the Oregon white oak woodland is to some extent a "fire type" then exclusion of fire by human influence was probably contributing to, if not largely responsible for, invasion by Douglas-fir. In this case the invasion is retrogression and reduction or loss of the herbaceous understorey is range depletion. If, contrariwise, invasion by and establishment of Pacific Douglas-fir ultimately results in a climatic climax Douglas-fir forest, then the Oregon white oak woodland is but a successional stage and therefore transitory forest range.
Both Kuchler (1964, 1966) and the U.S. Forest Service (Garrison et al., 1977, p. 33) interpreted Oregon white oak woodland as potential natural vegetation. These conclusions were accepted here.
It was concluded that invasion by Pacific Douglas-fir was likely due to unnatural (and imporper) exclusion of fire, a vegetational-determining part of the abiotic environment. Therefore invasion by Douglas-fir was a form of range retrogression and a brush problem. Good land stewardship and wise use management of this vegetation would include "burning the hell out of" this woodland range when perennial grasses, especially blue wildrye, were dormant. Or else the landowner should take ax in hand and spend the minimal effort required to chop out the non-sprounting Douglas-fir. Much of the range brush problem is at heart laziness and delayed response to what was initially a simple problem with an easy solution.
.Benton County, Oregon. June. Estival aspect. FRES No. 28 (Western Hardwoods Forest Ecosystem). K-22 (Oregon Oakwoods). SAF 233 (Oregon White Oak). Oregon White Oak Series in Oregonian Deciduous and Evergreen Forest biotic community or regional formation of Brown et al. (1998). Oregon white oak/blue wildrye association of Kagan et al. (2004).
17. Leader and leaves of Oregon white oak or Garry oak (Quercus garryana)- Benton County, Oregon. June.
17bbbb. Trunk of Oregon white oak- This Oregon or Garry oak in the Coast Range of northern California provided a textbook example of the bark pattern and color of its species. Most of the Oregon white oak farther north are hosts to so many species and such cover of mosses that their bark is seldom visible.
University of California Hopland Field Station, Mendocino County, California. June.
18. Yellow moss (Homalothecium fulgescens) on trunks of Oregon white oak- Yellow moss is one of the most common moss species on boles and branches of hardwood trees in many areas of the Coast Ranges. Note other bark covering species to left of yellow moss on the right bole.
Benton County, Oregon. June.
19. Yellow moss on trunk of Oregon white oak- Details of yellow moss on Garry oak. This is one of the most common epiphytes on angiosperm trees and shrubs throughout much of the Coast Ranges.
Benton County, Oregon. June.
20. Inflorescence of blue wildrye (Elymus glaucus)- Spike of blue wildrye. Benton County, Oregon. June.
21. Cluster or peafruit rose (Rosa pisocarpa)- This was one of two species of wild rose growing on an Oregon white oak woodland range in the Coast Ranges of northern Oregon. Photograph of flowering shoot was along Donner and Blitzen River, Harney County, Oregon. Close-up photograph of opened floral buds was in Garry oak woodland in Coast Ranges, Benton County, Oregon. The cluster arrangement of flowers (and later the rose hips) that was readily seen in the close-up slide was basis of the common name of this species.
22. Baldhip wildrose (Rosa gymnocarpa)- This wild rose was widely dispersed in the understorey of an Oregon white oak woodland community in the Coast Ranges of norther Oregon.
Benton County, Oregon. June.
"The Sierra Nevada and Cacade Mountains together constitute a single province but they are unlike in character, and the several sections of the Cascades alsodiffer among themselves" (Fenneman, 1931, p. 396). Fenneman (1931, ps. 396-441) divided "this great barrier" into the Sierra Nevada Section, Southern Cascade Mountains, Northern Cascade Mountains, and Middle Cascade Mountains. Franklin and Dyrness (1973, p. 6) followed a similar division of the Cascades into physiogrphic and geologic provinces and presented a generalized vegetation map that corresponded with their provinces and climatological parameters (Franklin and Dyrness, 1973, p. 45).
Some of the more important forest range types in the Cascade Mountains were presented in this section. Names, descriptions, and conclusions concerning this range vegetation followed that in Natural Vegetation of Oregon and Washington (Franklin and Dyrness, 1973), the definitive vegetation atlas and reference of these two states. Forest vegetation presented below was from the Western Cascades and High Cascades provinces designated by Franklin and Dyrness (1973, p. 6).
Forest range types of the Sierra Nevada were considered in a separate section as suggested by the above statement of Fenneman (1931, p. 396).
23. Landscape of Western Cascades- Various forest communities-- some seral; some climax-- developed in this part of the Middle Cascades Mountains section of Fenneman (1931, ps. 430-436) or, somewhat smaller and more specific as to vegetation, the Western Cascades of Franklin and Dyrness (1973, p.6). On zonal or regional scale vegetation seen here was part of the Tsuga heterophylla Zone of the latter authors. Much of it, however, was in late seral stages in Pacific Douglas-fir which had persisted into and continued to dominate the subclimax forests. Various disturbances-- notably wild fires, both natural and anthropogenic, and wind stroms-- late in the 1800s and early 1900s initiated secondary succession so that much of this largely re-developed vegetation approached its natural potential from perspective of species composition though not of community structure.
Within this general view were late seral (subclimax) western hemlock-western redcedar-Douglas-fir forests, Pacific silver fir-Douglas-fir forests, and subalpine fir-mountain hemlock forets (Kuchler units given below).
Among hardwoods there were many climax tree species, such as the bigleaf maple (Acer macrophyllum) growing here in the foreground, that were plentiful. There were also abundant populations of seral angiosperm tree and shrub species like bitter cherry (Prunus emarginata) and red alder (Alnus rubra). Climax conifers that in theory dominate the potential natural vegetation such as western hemlock (Tsuga heterophylla) and western redcedar (Thuja plicata) were generally of younger age classes that functioned as associates to the dominant subclimax Pacific Douglas-fir. Franklin and Dyrness (1973, ps. 70-87 passim) explained that Douglas-fir was often the sole dominant even in old-growth forests. This phenomenon was displayed in photogrphs that follow.
Herbaceous species were extremely varied in these three general climax forest communities, but were some forbs were common to most forests that were below subalpine. Ferns were some of the widely dispersed (distributed) forbs. Bracken fern (Pteridium aquilinium) and sword fern (Polystichum munitum) were most common (and in Coast Ranges forests as well).
Deschutes National Forest, Deschutes County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem), K-2 (Cedar-Hemlock-Douglas-fir Forest), SAF 229 (Pacific Douglas-fir) mostly but potential climax was SAF 227 (Western Redcedar-Western Hemlock) and FRES No. 23 (Fir-Spruce Forest Ecosystem), K-3 (Silver fir-Douglas-fir Forest), SAF (none applied), and K-4 (Subalpine Fir-Mountain Hemlock Forest), SAF 205 (Mountain Hemlock). Cascades- Western Cascades Montane Highlands Ecoregion, 4b (Pater et al., undated; Thorson et al., 2003).
24. Old-growth western hemlock-western redcedar-pacific Douglas-fir forest- Subclimax Pacific Douglas-fir was the dominant species and constituted almost all of the larger trees of this Western Cascades stand, but western hemlock and some (not much) western redcedar, the climax dominants, constituted most of the regeneration (except at the edge of this clearing that was in the foreground). There was also some regeneration of grand fir (Abies grandis).
Physiognomy of this forest type was portrayed in this subclimax old-growth stand.
Deschutes National Forest, Deschutes County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosysstem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 229 (Pacific Douglas-fir) with potential of SAF 227 (Western Redcedar-Western Hemlock). Cascade variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community or regional formation of Brown et a. (1998). Two associations presednted by Kagan et al. (2004) had developed in this forest vegetation: 1)western red cedar-western hemlock/dwarf Oregongrape and 2) western red cedar-western hemlock/beaked hazelnut/sword fern with dominant subclimax Douglas-fir in both associations. Cascades- Western Cascades Montane Highlands Ecoregion, 4b (Pater et al., undated; Thorson et al., 2003).
25. Passing of the guard- This closer-in and horizonal view of the old-growth western hemlock-western redcedar-Douglas-fir forest shown in preceding slide showed that the potential climax dominants, western hemlock and western redcedar, were succeeding the present dominant but subclimax Pacific Douglas-fir. At the edge of this forest (perimeter of a clearing from which the photograph was taken) there was regeneration of all three conifer species, but reproduction of Douglas-fir was restricted to this edge. Inside the forest reproduction of western hemlock (primarily) and western redcedar (limited) was overtaking that of Pacific Douglas-fir so that with advancing plant succession the latter species would be replaced by the two former specis, barring natural disturbances like fire or windstorms that could cause reinitiation of secondary plant succession. There was also some reproduction of grand fir that has greater tolerance (rated as Tolerant) than Pacific Douglas-fir (rated as Intermediate).
Deschutes National Forest, Deschutes County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 229 (Pacific Douglas-fir) on its way to SAF 227 (Western Redcedar-Western Hemlock).Cascade vriant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community of Brown et al. (1998). Two associations in this large-scale photo-plot were: 1) western red cedar-western hemlock/dwarf Oregongrape and 2) western red cedar-western hemlock/beaked hazelnut/sword fern (Kagan et al., 2004) with dominant subclimax Douglas-fir present in both. Cascades- Western Cascades Montane Highlands Ecoregion, 4b (Pater et al., undated; Thorson et al., 2003).
26. "Presidential Succession"- Old-growth specimens of Pacific Douglas-fir were slowly being "eased out" of dominance as rate of reproduction and production of young trees by western hemlock and, to much lesser degree, western redcedar were out-pacing that of subclimax Pacific Douglas-fir. Grand fir regeneration was also evident suggesting that this would be the main associate in the climax forest.
This vegetation was at outer edge of a large stand next to clearing. This was for purposes of securing a well-lite, detailed photograph but this unavoidably included more young Douglas-fir than in the interior of this subclimax forest. Largest tree was Pacific Douglas-fir with characteristic bark. Sapling to its left was also Douglas-fir (again there was more regeneration of this subclimax conifer along perimeter of forest). Pole-size tree to right of the large Douglas-fir was western hemlock. Smaller pole to the right (and farther back; deeper inside forest) was grand fir.
The broadleaf shrub at base of the large Pacific Dopuglas-fir was Douglas maple (Acer douglasii= A. glabrum var. douglasii). Other shrubs present in interior of this stand included Pacific rosebay (Rhodendron macrophyllum), California or beaked hazelnut (Corylus californica= C. cornuta var. californica), dull hollyberry or Oregon-grape or dwarf Oregongrape (Berberis nervosa= Mahonia nervosa), chinkapin or chinquapin (Castanopsis chrysophylla), rasberry (Rubus spp.), and several species of huckleberry, blueberry, or bilberry (Vaccinium spp.).
Herbaceous plants as to both species and cover. An herbaceous layer was present but discontinuous. A photographic sample of the herbaceous layer of a Douglas-fir-bigleaf maple forest was presented below.
Deschutes National Forest, Deschutes County Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 229 (Pacific Douglas-fir) with potential of SAF 227 (Western Redcedar-Western Hemlock). Cascade variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community or regional formation of Brown et al. (1998). Western red cedar-western hemlock/dwarf Oregongrape association of Kagan et al. (2004). Cascades- Western Cascades Montane Highlands Ecoregion, 4b (Pater et al., undated; Thorson et al., 2003).
27. Old-growth and second-growth- An old-growth stand of western hemlock-western redcedar-Pacific Douglas-fir towered above its regeneration on a roadcut in the Western Cascades province. Grand fir was more common than western redcedar but the latter species was included in the descriptive name of this stand to be consistent with cover types designations used by the Society of American Foresters. Reproduction on the disturbed area that was going through secondary succession was of all four of these major conifer species, but in the old-growth stand regeneration of the climax dominants, western hemlock and western redcedar, and of grand fir was greater than that of subclimax Douglas-fir as vegetation continued to develop toward the climax forest community.
There were some immense old-growth western hemlock (the large, overripe tree in right background was an example) and western redcedar, but these were far outnumbered (and outsized) by Pacific Douglas-fir.
There were also some Pacific silver fir (Abies amabilis) in this stand. This vegetation was high enough in elevation (a little above 4000 feet) that it also had sizable numbers of conifer species that were dominants in the Silver Fir-Douglas-fir Forest (Kuchler unit 3). This further complicated the successional status of Pacific Douglas-fir, but generally this conifer was seral. Layers of vegetation were well developed, but this was about as much due to various age classes of conifers as to presence of shrubs and herbaceous species.
Deschutes National Forest, Deschutes County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 229 (Pacific Douglas-fir) with potential of SAF 227 (Western Redcedar-Western Hemlock). Cascade variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community of Brown et al. (1998). Cascade variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community of Brown et al. (1q998). Over extent of forest range vegetation shown in this view there was some of both western red cedar-western hemlock/beaked hazelnut/sword fern and western red cedar-western hemlock/dwarf Oregongrape associations of Kagan et al. (2004) in both of which subclimax Douglas-fir was dominant. Cascades- Western Cascades Montane Highlands Ecoregion, 4b (Pater et al., undated; Thorson et al., 2003).
28. Changing forest- Vegetation is never static. Even climax plant communities at termination of succession are undergoing replacement of individuals of the climax species. Furthermore, any substantive disturbance can "move" the vegetation (climax as well as seral) back down the sere (ie. retrogression of the community). Even climax is not permanent. So too, later seral stages of vegetation may change (or appear to change) more slowly than preceding stages yet they are changing be it ever so slowly.
The forest vegetation shown here (an expanded horizonal view of the preceding "photo-plot") was an awe-inspiring old-growth stand of gargantuan Pacific Douglas-fir. These giant trees were seral (subclimax to be more accurate) and the "next generation" (ie. regeneration) was mostly of western hemlock, western redcedar, and grand fir (the first two being climax dominants for this forest cover type). The long-lived individuals of the seral Douglas-fir had persisted into the early years of the climax western hemlock-western redcedar-Douglas-fir forest (Kuchler unit 2) in the Western Cascade province.
An old highway right-of-way was in the foreground of this photograph and on this "new land" (= denuded, disturbed soil) regeneration was of all four coniferous species, but inside Pacific Douglas-fir was loosing the dominance race.
Deschutes National Forest, Deschutes County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Dougla-fir Forest). SAF 229 Pacific Douglas-fir) with successional potential for SAF 227 (Western Redcedar-Western Hemlock). Cascade variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community or regional formation of Brown et al. (1998). Two Kagan et al. (2004) associations were represented with persistent subclimax Douglas-fir dominant in both: 1) western red cedar-western hemlock/dwarf Oregongrape and 2) western red cedar-western hemlock/beaked hazelnut/sword fern. Cascades- Western Cascades Montane Highlands Ecoregion, 4b (Pater et al., undated; Thorson et al., 2003).
29. The dynamics of forest vegetation (changing of dominance up close)- It was discussed repeatedly above that Pacific Douglas-fir is subclimax on most forest types and sites even where it is dominant (even where it is dominant into climax). That condition or phenomenon was shown in this small sample of of forest vegetation from the Western Cascades province.
A large Douglas-fir (largest tree; big trunk in left foreground) was the reigning monarch, but it was not a hereditary monarchy. The barbarians were by the bark of the old monarch and scaling the successional wall. Western redcedar (represented by a large seedling in center foreground) and western hemlock (large saplings and small poles surrounding the Douglas-fir) were succceeding Douglas-fir as dominant species of the climax Cedar-Hemlock-Douglas-fir Forest (K-2), western redcedar-western hemlock (SAF 227).
The low shrubs in center foreground were a Rubus species. Sword fern was also conspicuous. Other herbaceous species included Oregon sorrel (Oxalis oregana), tall bugbane (Cumicifuga elata), and giant or western wake-robin (Trillium chloropetalum= T. sessile).Dull hollygrape or dull Oregon-grape was the most common shrub species in the largely herbaceous understorey. There were no grasses or grasslike plants in understorey.
Farther downslope (right background) bigleaf maple was well-established.
Willamette National Forest, Linn County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedr-Hemlock-Douglas-fir Forest). SAF 227 (Western Redcedar-Western Hemlock) is climax. Cascade variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community of Brown et al. (1998). Western red cedar-western hemlock/dwarf Oregongrape association (Kagan et al., 2004). Cascades- Western Cascades Montane Highlands Ecoregion, 4b (Pater et al., undated; Thorson et al., 2003).
30. The bottom of the bottom (floor of a bottomland Western Cascade forest)- The lowest layer of vegetation in a Pacific Douglas-fir-bigleaf maple riparian forest on a terrace of the Santiam River was composed of herbs and low-growing shrubs. Common species in this diverse understorey included Oregon sorrel, giant wake-robbin, star-flowered false Solomon's seal (Smilacina stellata), and dull hollygrape or dull Oregon-grape.
Black cottonwood (Populus trichocarpa) appeared to be the associate species. While there were enough large Douglas-fir that this species dominated the canopy there were also large individuals of western hemlock western redcedar that clearly identified this as a potential western redcedar-western hemlock-Douglas-fir forest. There were numerous small groves of bigleaf maple indicating that this species was a forest dominant (at least locally).
Linn County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). It was likely that frequent flooding of the Santiam River created enough exposed mineral soil for Douglas-fir seedbeds that this species was likely to remain co-dominant with bigleaf maple. As such this was designated SAF 229 (Pacific Dougls-fir) rather than the climax 227 (Western Redcedar-Western Hemlock). Cascades- Western Cascades Lowlands and Valleys Ecoregion, 4a (Pater et al., undated; Thorson et al., 2003).
31.Riparian forest of bigleaf maple- This section of riparian corridor along the Santiam River in the Western Cascades was a bigleaf maple community. Some of the trees were relatively small, but this consociation of a hardwood species that is rated as Tolerant (Wenger, 1984, p.7) was interpreted as the climax vegetation for this bottomland site. Much of the understorey layer was a younger age class of bigleaf maple.
Linn County, Oregon. June. No FRES or Kuchler unit for vegetation at this small scale. Bigleaf maple-Douglas-fir/beaked hazelnut/ Pacific waterleaf (Hydrophyllum tenuipes) association of Kagan et al. (2004). Cascades- Western Cascades Lowlands and Valleys Ecoregion, 4a (Pater et al., undated; Thorson et al., 2003).
32. Bigleaf maple forest- A climax bigleaf maple community had developed on this riparian zone of the Santiam River in the Western Cascades province. This species has a tolerance rating of Tolerant (Wenger, 1984, p. 7). Based on this "photo-plot" that reputation was well-deserved. Viewers should note that tree regenertaion was exclusively bigleaf maple. In fact, the understorey was dominated by a younger age class of bigleaf maple. This species was reproducing in its own shade.
Franklin and Dyrness (1973, p. 126) remarked that bigleaf maple was a common riparian community in some areas but that it was "not particularily charcteristic".
Linn County, Oregon. June. No FRES or Kuchler because these sources did not map vegetation at the riparian scale. Bigleaf maple-Douglas-fir/beaked hazelnut/Pacific waterleaf association of Kangan et al. (2004). Cascades- Western Cascades Lowlands and Valleys Ecoregion, 4a (Pater et al., undated; Thorson et al., 2003).
33. Pacific Douglas-fir, the pioneer- Large, old-growth Douglas-fir so often persist into the climax that it would be easy to forget or overlook the fact that this mighty conifer can often be a pioneer species. Pacific Douglas-fir readily invades denuded sites, especially those with scarified soil (exposed mineral soil), and becomes established early in secondary succession. This is why clearcutting and seed tree methods are widely used to regenerate the next crop of Douglas-fir.
That principle was evident in this photograph. A roadcut had been pioneered by Douglas-fir seedlings that invaded the fresh site naturally from seed shed by upslope trees. These little fellows are likely to grow into mature trees and remain in developing forest vegetation even if and when climax conifers like western hemlock or grand fir become adult trees.
This was the start of a Douglas-fir forest.
Willamette National Forest, Linn County, Oregon. FRES No. 20 (Dougls-fir Forest Ecosystem). Potential natural vegetation unknown so now Kuchler unit presented. SAF 229 (Pacific Douglas-fir). Cascades- Western Cascades Montane Highlands Ecoregion, 4b (Pater et al., undated; Thorson et al., 2003).
34. Western hemlock climax forest- This vegetation was a representative composite view of the western hemlock-chinkapin (Tsuga heterophylla/Castanopsis chyrsophylla) association described by Franklin and Dyrness (1973, p. 76 [mostly]-81 passim). This is the driest of the forest associations within the Tsuga heterophylla Zone of the Western and High Cascades provinces to be dominated by western hemlock. In an array or spectrum (zonation) of these forest associations the Pseudotsuga menziesii/Holodiscus discolor association is the driest (least mesic) and the Tsuga heterophylla/Polystichum munitum-Oxalis oregana association is the most mesic. Much, if not most, of the latter has been converted into industrial Pacific Douglas-fir forests.
An example of the Douglas fir-oceanspray association from the Olympic Peninsula province was presented in the Olympic Peninsula portion of the section, Pacific Northwest (Forests). That example of forest vegetation can be compared with this western hemlock-chinkapin association and the western hemlock-California rosebay association presented below to visualize the gradient and zonation of habitats within the Tsuga heterophylla Zone.
This exterior perspective of the western hemlock-chinkapin association (climax vegetation for this forest site) in the High Cascades showed the physiognomy of this subtype of the western hemlock cover type (SAF 230) and of the range type overall. This was also evident in the next association shown below. The shrub layer of this (and the next) understorey gave a two-layered general appearnce, but there was also at least one (arguably, two) other layers of vegetation, including a species-ric herb layer dominated by forbs.
From standpoint of range management this was a browse range with chinkapin being the major borwse species. Dull hollygrape was also a common shrub, and its substantial cover of lower stature would allow the shrub part of this forest to be interpreted as consisting of two layers.
Many (probably most) of the larger trees in this vegetation were Pacific Douglas-fir that had become established early in succession and lived into the climax stage. These larger Douglas-fir became established on the site at earlier seral stages and lived throughout progression of the sere. In this second-growth forest Douglas-fir may have invaded the cut-over land at the initial or pioneer stage of secondary succession as shown in the immediately preceding slide.
Warm Springs Indian Reservation, Wasco County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 230 (Western Hemlock) most descriptively as western reddcedar was a minor component. Students should have realized at this point that the spatial scales of the FRES and Kuchler designations are frequently too large to distinguish units of vegetation at the scale of forest (and rangeland) cover types. Distinction of this climax forest vegetation from that of the preceding climax forest was at cover type level (ie. SAF 229 vs. SAF 230). Cascade variant of Western Hemlock Series in Coastal Conifer Forest biotic community of Brown et al. (1998). Western hemlock/golden chinquapin assocition of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
35. Interior of a western hemlock-chinkapin climax forest- Inside the same "plot" or stand of forest vegetation of the Tsuga heterophylla-Castanopsis chysophylla association shown "from the outside" in the preceding "photographic quadrant".
The largest tree (trunk in left foreground) was a Pacific Douglas-fir that had lived from early in the sere into the present climax state. All of the young trees were western hemlock or, less abundant, western redcedar. Chinkapin grew throughout the understorey sometime being the sole species in the understorey, but more commonly having dull Oregon-grape or dull hollygrape and numerous forbs as associated plants. Bracken fern was "never far away". Various kinds of huckleberry, blueberry, or bilbery (Vaccinium spp.) were also present.
Various of these understorey forest species were introduced below. All of these examples were photographed in the same forest stand shown in this and the preceding slide.
One of various kinds (cover types and subtypes) of forests in the High Cascades physiographic and geologic province. The Middle Cascades section of the Sierra-Cascade province of Fenneman (1931, ps. 430-436).
Warm Springs Indian Reservation, Wasco County, Oregon. June. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 230 (Western Hemlock) because western redcedar was not co-dominant with western hemlock. The latter was the potential dominant tree of this association (Franklin and Dyrness, 1973. p. 76).Cascade variant of Western Hemlock Series in Oregonian Coasal Conifer Forest biotic community of Brown et al. (1998). Western hemlock/golden chinquapin association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
36. Floor of the forest- Two layers of the understorey of the western hemlock-chinkpin climax forest. Chinkapin (also chinquapin) was the shrub with dark green leaves along left margin. Chinkapin dominated the shrub layer. Dull hollygrape, a lower-growing shrub, often formed a second shrub layer. The herbaceous layer (rest of photograph to right of chinkapin) consisted of several species of forest forbs several of which follow.
Warm Springs Indian Reservation, Wasco County, Oregon. FRES No. 20 (Douglas-fir Forest Formation). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 230 (Western Hemlock). Cascade variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community of Brown et al. (1998). Western hemlock/golden chinquapin association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
37. Bigleaf avens (Geum macrophyllum)- This herbaceous member of the rose family was a local dominant forb in the western hemlock-chinkapin climax forest.
Warm Springs Indian Reservation. Wasco County, Oregon. June.
38. Woods strawberry (Fragaria vesca var. crinita= F. crinita)- Like the preceding forb the wild strawberry is a member of the rose family. In fact both are members of the same subfamily of Rosaceae, the Rosoideae. This is one of at least three Fragaria species that live in the big woods of the Pacific Northwest. Western hemlock-chinkapin forest in High Cascades province.
Warm Springs Indian Reservation, Wasco County, Oregon. June..
39. Dull hollygrape or Dull Oregon-grape (Berberis nervosa= Mahonia nervosa)- The barberry family is a small one genus-wise, but some of its species are important due to their dominance role and widespread occurrence. Dull hollygrape is an example. Dull Oregon-grape is a major low-layer understorey, evergreen shrub in several forest types including the various Douglas-fir-dominated forests, western white pine forests, and western hemlock-dominated forests.This specimen was growing in the same western hemlock-chinkapin climax forest shown above.
Note fruit near base of this individual.
This and various other species in the Berberis-Mahonia group have long been used as ornamentals in landscaping.
Warm Springs Indian Reservation. Wasco County, Oregon.
40.Three-leaf windflower or three-leaf anemone (Anemone deltoidea)- This member of the buttercup family was common and locally very abundant on the floor of a western hemlock-chinkapin forest in the High Cascades province. Throughout much of the Western Range Region-- on both forest and rangeland-- various members of the Ranunculaceae are important and often showy forbs.
Warm Springs Indian Reservation, Wasco County, Oregon. June.
41.Queen's cup, bride's bonnet, or blue-bead (Clintonia uniflora)- In the Pacific Northwest the beadlily genus includes one of many species in the diverse lily family. This particular species is one of the most showy and readily recognized forbs ("wild flowers") affilitated with the the vast and varied coniferous forest formation in northwestern North America. In flower or fruit this beadlily is a real beauty and a delightful treat to those souls fond of Nature's granduer.
Western hemlock-chinkapin climax forest in Western Cascades province (samd stand as all other species in this portion).
Warm Springs Indian Reservation, Wasco County, Oregon.
42. American or northern starflower (Trientalis borealis)- Northern starflower belongs to the primrose family and is another of the showy forbs of the northwestern coniferous forest. This one was another of the nextdoor-neighbor species introduced to viewers from the stand of western hemlock-chinkapin forest in the High Cascades physiographic and geologic province.
Warm Springs Indian Reservation, Wasco County, Oregon. June.
43. Spotted western corralroot (Corallorhiza maculata var. maculata)- This saprophytic orchid seemed appropriate to be the last of several "forbaceous" "characters" that represented the western hemlock-chinkapin climax forest. Unlike other understorey plants this botanical prize of wild flower connoisseurs is a saprophyte, a plant (plant species) that is incapable of manufacturing its own photosynthate and instead is dependent on dead, decaying organic matter as sources for its nourishment. In the ecosystem concept this is the biotic (organism) group designated as decomposers or reducers. Although microorganisms and fungi are the more common and widely distributed decomposers some vascular plants are reducers. This beautiful orchicd was one such species. It was included here as an example of both: 1) a forb species that ws a member of the understorey of the western hemlock-chinkapin forest and 2) a decomposer of the natural ecosystem.
This individual was growing in the same western hemlock forest stand as the other plant species included in this portion.
Warm Springs Indian Reservation, Wasco County, Oregon.
44. Western hemlock-chinkapin climax forest- This forest stand was "almost identical" to the one portrayed immediately above except that the herbaceous understorey species were limited being largely restricted to beargrass (Xerophyllum tenax). The driest sites of the western hemlock-chinkapin association are characterized by populations of this conspicuous forb (Franklin and Dyrness, 1973, ps. 74, 76, 79, 80, 82).
The forest community growing on this shallow stoney site in the High Cascades province should be contrasted with the form of the western hemlock-chinkapin association that developed on the more moist, deeper-soil site shown immediately above and the western hemlock-Pacific rosebay association shown immediately below.
Pacific Douglas-fir comprised much of the older-age class stocking of this stand, but western hemlock was succeeding it with most regeneration limited to the latter conifer species (the potential climax dominant).
Warn Springs Indian Reservation, Wasco County, Oregon. FRES No. 20 ( Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 230 (Western Hemlock). Cascade variant of Oregonian Coastal Conifer Forest biotic community of Brown et al. (1998). Western hemlock/golden chinquapin association of Kagan et al. (2004), but with beargrass sometimes locally dominant so as to have the western hemlock/beargrass association at local (microsite) scale. Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
45. Postcards of High Cascades western hemlock-dominated forest- Two picture-perfect views of the western hemlock-Pacific rosebay-dull hollygrape (Tsuga heterophylla/Rhodendron macrophyllum/Berberis nervosa) association in the Tsuga heterophylla Zone (Franklin and Dyrness, 1973, ps.73-88 passim). This climax stand in the Central Cascades section (Fenneman, 1931, ps. 430-436) still had considerable canopy cover of Pacific Douglas-fir, but there were also several old-growth specimens of western hemlock and, though fewer, of western redcedar. Evidence of eventual dominance by the Very Tolerant western hemlock was the overwhelming regeneration of this species and not Douglas-fir.
The western hemlock-Pacific (California) rosebay-dull hollygrape association was interpreted by Franklin and Dyrness (1973, p. 73-78) as being the fourth in an array of moisture gradients from the driest Douglas-fir-oceanspray to the wettest western hemlock-sword fern-Oregon sorrel associations. Species relations did not always seem consistent across this moisture gradient. Beargrass was viewed by Franklin and Dyrness, 1973, ps. 74, 76, 79, 80, 82) as characteristic of the relatively dry western hemlock-chinkapin association yet beargrass was also common, even locally dominant, in the more mesic western hemlock-Pacific rosebay-dull hollygrape association which in case of this stand supported numerous western redcedar, a species with generally greater soil moisture requirements than western hemlock and Douglas-fir (Harlow et al., 1979, p. 196; Wenger, 1984, p. 6; Franklin and Dyrness, 1973, p. 81-82). Franklin and Dyrness (1973, p. 82) concluded with this qualifying statement: "There are many variations of the general community pattern throughout the Tsuga heterophylla Zone". These investigators interpreted the western hemlock-Pacific rosebay-dull hollygrape as typifying "the climatic climax for the Tsuga heterophylla Zone" (Franklin and Dyrness, 1973, p.77). Rhodendron macrophyllum is known variously as California rosebay, Pacific rosebay, California rhodendron, and Pacific rhodendron.
Mount Hood National Forest, Wasco County, Oregon. FRES No. 20 (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 227 (Western Redcedar-Western Hemlock) as distinguished from the preceding SAF 230 (Western Hemlock) by more western redcedar (local co-dominance) on a more mesic habitat. Cascade variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community of Brown et al. (1998). Western hemlock/western rhodendron (R. macrophyllum) association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003). Note: use of "western rhodendron" in association was a poor choice of common name and, in fact, as close to improper use of a common as it gets because Rhodendron occidentale grows in Oregon and this is obviously the "real" western rhodendron.
46. Local stand of young western hemlock and associated species in High Cascades- Detail "quadrant" of local vegetation in western hemlock-Pacific rosebay-dull hollygrape association. On this slightly disturbed microsite western hemlock, beargrass, and resprouts of trailing snowberry, red huckleberry, California or blue huckleberry, and chinkapin comprised the plant community. Hollygrape was most frequently dominant under shade. Rosebay was most abundant on openings and older clearings were light had not been limiting durin establishment.
With obvious exception of beargrass there were very few forbs and no other monocots in understorey. Once again with late seral and climax coniferous forests in the Cascade Mountains forest range was browse range.
Mount Hood National Forest, Wasco County, Oregon. FRES No. (Douglas-fir Forest Ecosystem). K-2 (Cedar-Hemlock-Douglas-fir Forest). SAF 230 (Western Hemlock) at scale of local stand; SAF 227 (Western Redcedar-Western Hemlock) overall. Cascde variant of Western Hemlock Series in Oregonian Coastal Conifer Forest biotic community of Brown et al. (1998). Western hemlock/western rhodendron (R. macrophyllum)-evergreen huckleberry (Vaccinium ovatum) association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
47. Pacific silver fir-dominated forest- General view of climax Pacific silver fir forest in the High Cascadaes of northern Oregon. This climax forest community was a representative sample of the Abies amabilis-Tsuga mertensiana/Vaccinium membranaceum association described by Franklin and Dyrness (1973, ps. 96-98, 103-105). It was underscored at this point that these authors placed and described this association in both the Tsuga mertensiana and the Abies amabilis Zones with switching of the two co-dominants back-and-forth even within the same (regardless of the) Zone (Franklin and Dyrness, 1973, ps. 98, 104-105).
Two adult (mature) trees in foreground were Pacific silver fir with the flaky bark characteristic of old trees (though bark of these two individual boles had a brownish cast absent from most other contemporaries). Reproduction in exposed mineral soil in foreground was all silver fir. Numerous mountain hemlock were visible in a more interior part of the forest (background).
The understorey (not visible behind the dense cover of silver fir seedlings in foreground; this understorey was shown in the third slide immediately below) was a consociation of black or thinleaf huckleberry or blueberry (Vacinnium membranaceum) with some patches of fool's huckleberry or rusty leaf (Menziesia ferruginea). Vanillaleaf or deer's foot (Achyls triphylla) was the major forb. Beargrass was present, but in trace amounts and with irregular distribution (random dispersion). Idaho fescue (Festuca idahoensis) was rare in forest gaps.
Mount Hood National Forest, Hood River County, Oregon. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock Forest), closed forest subzone: this unit was not really the most appropriate, but K-3 (Silver Fir-Douglas-fir) was even less descriptive given rarity of Douglas-fir and near co-dominance of Pacific silver fir and mountain hemlock in the stand. On basis of species dominance on the sere during plant succession (rather than on species dominance at climax), K-3 (Silver Fir-Douglas-fir Forest) would be the appropriate unit because Pacific Douglas-fir often pioneers the sere and may continue to dominate successional vegetation for four to five hundred years post-denudation and initial invasion (Franklin and Fyrness, 1973, ps. 98-99).
It was explained below where the High Cascades mountain hemlock forest type was presented that this area appeared to be a transition (ecotone may, or may not, apply) between the Abies amabilis and Tsuga mertensiana Zones. Recall from the first paragraph of this caption that relative dominance of these co-dominant conifer varied across the two Zones. There was no SAF designation for Pacific silver fir so "officially" a Pacific silver fir variant of SAF 205 (Mountain Hemlock). Pacific Silver Fir Series in Cascade-Sierran Montane Conifer Forest biotic community or regioal formation of Brown et al. (1998). Pacific silver fir/thin-leaved blueberry/vanillaleaf association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
48. Dense stand of Pacific silver fir- This two-age classes stand of Pacific silver fir was representative of pure stands, "stands in which at least 80 percent of the trees in the main crown canopy are of a single species" ((Hawley, 1950), of this species that developed at midelevations in the High Cascades of northern Oregon. Seedling regeneration was restricted to forest perimeters and gaps as was understorey development. (This was more evident in the next, immediately succeeding, slide.) The forb in the close foreground was deer's foot or vanillaleaf. Low shrub in understorey at edge of stand was black huckleberry.
There were some widely scattered mountain hemlock deeper in this stand, but overall this forest was a pure stand of Pacific silver fir.
Mount Hood National Forest, Hood River County, Oregon. June. FRES No. 23 (Spruce-Fir Forest Ecosystem). No Kuchler unit or SAF cover type designation seemed appropriate. Discussion of these vegetation units under preceding caption was even more fitting here. Pacific Silver Fir Series in Cascade-Sierran Montane Conifer Forest biotic community of Brown et al. (1998). Pacific silver fir/vanillaleaf association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
49. Pure stand of Pacific silver fir in Oregon High Cascades- Another view of the two-age classes stand of Pacific silver fir presented in the preceding slide. The forest floor in interior of the stand was devoid of any understorey. Fallen trunks and crowns were more common than herbs, shrubs, or silver fir seedlings. A lower layer of vegetation comprised of black huckleberry and here-and-there patches of deer's foor or vanillaleaf was limitead to forest edges and gaps as was silver fir reproduction.
Mount Hood National Forest, Hood River County, Oregon. June. FRES No. 23 (Spruce-Fir Forest Ecosystem). None of the Kuchler units or SAF cover type designations "fit" (aptly descriped) to this pure stand of Pacific silver fir. See discussion in caption for first photograph of the Pacific silvre fir forest series. Pacific Silver Fir Series in Cascade-Sierran Montane Forest biotic community of Brown et al. (1998). Pacific silver fir/vanillaleaf association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
50. Climax Pacific silver fir forest- Interior of a Pacific silver fir forest in High Cascades of northern Oregon. This was an uneven-aged stand of Pacific silver fir. An uneven-aged stand is one "in which there are considerable differences in age of trees and in which three or more age classes are repersented " (Hawley, 1950) or "a stand with trees of three or more distinct age classes, either intimiately mixed or in small groups" (Helms, 1998). In fact, this was more likely an all-aged stand, "a stand in which theoretically trees of all ages up to and including those of the felling age are found" (Hawley, 1950) or "a stand with trees of all or almost all age classes, including those of exploitable age" (Helms, 1998).
There were a few, widely scattered mountain hemlock in this forest, but this was both an uneven-aged or all-aged stand and a pure stand. Presence of all, or nearly all, age classes including-- most revealingly-- seedling illustrated the Tolerant tolerance rating (Wenger, 1984, p. 3) of this "the major climax species" of the Abies amabilis zone in which (Franklin and Dyrness, 1973, p. 98). Franklin and Dyrness (1973, p. 99) specified that evaluations of stand structure showed that mountain hemlock was "a minor climax species" in the Pacific silver fir zone so that on wet sites the Very Tolerant mountain hemlock (Wenger, 1984, p. 3) is replaced by silver fir. This is probably because this hemlock seedlings cannot survive under the accumulations of snow characateristic of this zone whereas seedlings of silver fir can. Furthermore, regenertion of mountain hemlock is usually restricted to accumulations of organic matter such as nurse logs and other large pieces of rotting debris.
Bark on trunks seen here was characteristic of that on mature and old trees. Bark on the largest bole in left foreground was the textbook grey, flaking, chip-covered bark on very old Pacific silver fir.
The understorey was a consociation of black or thinleaf huckleberry or blueberry. There were traces of beargrass, fool's huckleberry, and vanillaleaf. This was definitely browse range.
Mount Hood National Forest, Hood River County, Oregon. June. FRES No. 23 (Spruce-Fir Forest Ecosystem). The only Kuchler units that come close to "matching" (ie. a "close fit") a pure stand of Pacific silver fir were K-3 (Silver Fir-Douglas-fir Forest) or K-4 (Fir-Hemlock Forest). Readers were referred back to discussion of this problem in first caption for the Pacific silver fir forest series. No SAF cover type designation for Pacific silver fir so "officially" this was Pacific silver fir-variant of SAF 205 (Mountain Hemlock) given the species affinities of the adjoining Abies amabilis and Tsuga mertensiana Zones (Franklin and Dyrness, 1973, 93-108 passim). Abies amabilis-Tsuga mertensiana/Vaccinium membranaceum association given by Franklin and Dyrness (1973, p. 98, 104-105) where Tsuga mertensiana is local co-dominant or even dominant depending on forest site.
Pacific Silver Fir Series in Cascade-Sierran Montane Conifer Forest biotic community of Brown et al. (1998). Pacific silver fir/thin-leaved blueberry/vanillaleaf association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
51. Vanillleaf or deer's foot (Achlys triphylla)- This barberry familymember was the local dominant (the only) forb in a pure stand of Pacific silver fir, and the species was restricted to forest edges and gaps. Mount Hood National Foret, Hood River County, Oregon June..
52. Mountain hemlock (Tsuga mertensiana) forest- Physiognomy of a mountain hemlock-dominated forest in the High Cascades geologic and physiographic province. "The Tsuga mertensiana Zone is the highest forested zone along the western slopes and crest of the Cascade Range and in the Olympic and Klamath Mountains" (Franklin and Dyrness, 1973, p. 101). With that for introduction, students were referred once more to the "vegetation bible" of Franklin and Dyrness (1973, p. 101-109) for coverage of forest communities in this mountain region.
This mountain hemlock-dominated stand was far from from a pure stand, one "composed of of essentially a single species" (Helms, 1998). Western hemlock, grand fir, Pacific silver fir (Abies amabilis) were also common. There were a few omnipresent Pacific Douglas-fir. Western redcedar was absent. Regeneration of conifers (young saplings on exposed soil in foreground) was primarily Pacific silver fir. Hemlock (both Tsuga species) survival as seedlings requires abundant organic matter such as that provided by nurse logs.
The understory vegetation of the mountain hemlock forest was limited in both species and structure with the main shrub being black or thinleaf huckleberry (Vaccinium membranaceum). There was less cover of grouseberry or whortleberry (V. scoparium= V. erythrococcum= V. myrtillus var. microphyllum) and much less of the uncommon common pipsissewa (Chimphila umbellata). Beargrass was widespread along the perimeter of forest openings. An example of a diverse herbaceous-woody understorey in openings among the closed canopy forest was included below.
Mount Hood National Forest, Hood River County, Oregon. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock Forest). Note: this is the lower subzone closed forest of the Tsuga mertensiana Zone described by Franklin and Dyrness (1973, ps. 101-102). Variant of SAF 205 (Mountain Hemlock). Mountain Hemlock Series in Cascade-Sierran Subalpine Conifer Forest biotic community of Brown et al. (1998). Mountain hemlock/thin-leaved blueberry/beargrass association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
53.Mountain hemlock on a mountain slope- A climax mountain hemlock forest had developed on this west slope of the Oregon Cascade Range. This vegetation formed the highest elevation forest in the High Cascades province. Mountain hemlock was the dominant but Pacific silver fir, western hemlock, grand fir, and even a few Douglas-fir contributed to the stocking. Regeneration on the mineral soil of the foreground was mostly silver fir with some grand fir.
Both hemlock species usually require organic matter for seedling survival. Mountain hemlock and Pacific silver fir are climax species and they often grow together, but regeneration on mineral soil is limited to the latter species (Franklin and Dyrness, 1973, ps. 99-100). That situation was evident in this and the immediately preceding photograph where young trees (splings in the first photograph and large seedlings in the second) on road cuts were Pacific silver fir. In the second slide two large mountain hemlocks (center edge of mature trees) had portions of main roots above ground surface line with the spreading pattern indicative of establishment on nurse logs.
The lower-elevation, closed forest form (subzone) of mountain hemlock zone intergrades with the Pacific silver fir zone that is below it in elevation. Some ecologists have interpreted the closed forest of mountain hemlock to be a part of phase of the Pacific silver fir forest (Franklin and Dyrness, 1973, p. 102).
This closed stand and the one in the preceding slide had a relatively sparse understorey. The lower level of vegetation was limited primarily to black huckleberry. An example of a well-developed understorey present in openings within the closed canopy forest was shown immediately below.
Mount Hood National Forest, Hood River County, Oregon. June FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock Forest). Note: this is the lower subzone closed forest of the Tsuga mertensiana Zone described by Franklin and Dyrness (1973, p. 101-102). SAF 205 (Mountain Hemlock). Mountain Hemlock Series in Cascade-Sierran Subalpine Conifer Forest biotic community of Brown et al. (1998). Mountain hemlock/thin-leaved blueberry/beargrass association of Kagan et al. (2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
54.Transition Pacific silver fir-mountain hemlock forest with understorey vegetation- This old-growth forest in the High Cascades of northern Oregon appeared to be an ecotone between Pacific silver fir forest and the lower elevation subzone of mountain hemlock forest with a mixture of conifer species common to both types including western hemlock, Pacific Dougals-fir, grand fir, and a few noble fir (Abies procera).
There was a fairly large opening around a seep or spring in this forest vegetation. Species compostion of this opening was extraordinarily rich in comparison to the more usual impoverished understorey of these two cover types. Species included black huckleberry, whortleberry, raspberry (Rubus spp.), one-sided wintergreen or sidebells pyrola (Pyrola secunda= Orthiia secunda), common pibsissewa, beargrass, and fool's huckleberry or rustyleaf (Menziesia ferruginea). The majority of these species were in the heath or heather family (Ericaceae). Also growing along the seep were large specimens of swamp latern or western skunk cabbage (Lysichitum americanum). Grasses and grasslike plants were almost absent except for occasional individuals of Idaho fescue (Festuca idahoensis) and pinegrass (Calamagrostis rubescens). Apparently coniferous species of the Cascades require well-drained soils. They were absent from this drainage area except along the edges of it. Franklin and Dyrness (1973, p. 106) gave a brief description of vegetation around seeps in mountain hemlock forests.
Vegetation in this transition between Pacific silver fir and mountain hemlock was perhaps not so much an ecotone as a stage in forest community development. In the mountain hemlolock zone Pacific silver fir is often a climax associate to mountain hemlock and vice versa for mountain hemlock in the Pacific silver fir zone such that distinction between these two forest cover types is one of relative species predominance and, by its very nature, rather arbitrary. Matters are further complicated by the mosaic of these forest range types due to slope, aspect, soils, sooil seed bank, fire regime, and past human impaact.
Mount Hood National Forest, Hood River County, Oregon. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock Forest). Variant of SAF 205 (Mountain Hemlock). There is no SAF designation for Pacific silver fir-dominated forest. Instead Pacific silver fir (and noble fir) were included with SAF 205. The view of some ecologists that the closed forest form or subzone of mountain hemlock is actually part of the Pacific silver fir zone (Franklin and Dyrness, 1973, 102) was cited in the previous caption and bore repeating at this point. Transition or "hybrid" between Pacific Silver Series in Cascade-Sierran Montane Conifer Forest biotic community and Mountain Hemlock Series in Cascade-Sierran Subalpine Conifer Forest biotic community of Brown et al. (1998). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
55. Mountain hemlock stand- Understorey and regeneration of mountain hemlock-dominated climax forest of the closed forest subzone form of this forest cover type. This stand of forest vegetation in the High Cascades was dominated by mountain hemlock with Pacific silver fir as the associate conifer. There was regenertion of both species, but with reproduction of mountain hemlock restricted to the interior where there was adequate organic matter for survival of its seedlings.
The understorey was dominated by ericaceous (of the Ericaceae, the heather or heath family) shrubs and subshrubs, especially Vaccinium species and rustyleaf or fool's huckleaf. There were rare individuals of the cespitose Idaho fescue that appeared to be of little consequence in the forest community except to add diversity. The species was noted here only as a vegetation "benchmark" to rangemen.
Mount Hood National Forest, Hood River County, Oregon. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock Forest), closed forest subzone of Mountain Hemlock. Variant of SAF 205 (Mountain Hemlock). Mountain Hemlock Series in Cascade-Sierran Subalpine Conifer Forest biotic community of Brown et al. (1998). In this stand there were locally two small examples of both: 1) mountain hemlock/thin-leaved blueberry/beargrass and 2) mountain hemlock/fool's huckleberry associations (Kagan et al., 2004). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
56. When plants tell tales but don't play by the rules- This eye-catching, old-growth stand of High Cascades forest vegetation illustrated a plant community that defied a vegetation mappers pen (perhaps even imagination). This forest range vegetation had developed in a deep cove where Pacific silver fir and mountain hemlock were dominant species, but where these Tolerant and Very Tolerant, respectively, climax species had been joined by grand fir (also Tolerant) and (of course) even a few Pacific Douglas-fir. Some of the largest trees, including those like Jack-who-fell-down-and-broke-his-crown, were grand fir.
The species composition of the conifer component was such as to bely typing of vegetation. The forest (and forest range) communitiy was as much a hodgepodge as the jaxed fallen timbers. This forest "sample" would probaly be most accurately described as the Abies amabilis-Tsuga mertensiana/Vaccinium membranaceum association (Franklin and Dyrness, 1973, p. 104-105) of the Tsuga mertensiana Zone.
The understorey was overwhelmingly woody with black huckleberry and grouseberry as dominants. most common species. One-sided wintergreen and fool's huckleberry or rustyleaf were present, even locally abundant.
The structure, age classes, and, especially, physical condition of individual trees of the forest vegetation in these "photo-quadrants" were typical and indicative of old-growth forest. This was not only a virgin forest, it was a virgin forest that had reached climax successionally and old-age physiologically. Trees with broken off and fallen crowns (nothing but standing logs) and downed timber (often criss-crossed upon itself) were textbook signs of ancient trees that were long past their "natural prime".
This was an example of sawtimber, "trees or logs cut from trees with minimum diameter and length and with stem quality suitable for conversion to lumber" (Helms, 1998), that had long ago become overmature. Overmature: "a tree or even-aged stand that has reached that stage of development when it is declining in vigor and health and reaching the end of its natural life span" or "a tree or even-aged stand that has begun to lessen in commercial value because of size, age, decay, or other factors", but "note the term has little applicability to uneven-aged stands, which consist of trees of diverse ages and stages of development" (Helms, 1998).
Regeneration had occurred (was occurring) in this stand, but for practical purposes it was an even-aged stand, "a stand of trees composed of a single age class in which the range of tree ages is usually [within] 20% of rotation" where rotation is "in even-aged stands, the period between regeneratioin establishment and final cutting" (Helms, 1998).
Mount Hood National Forest, Hood River County, Oregon. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-2 (Fir-Hemlock). Variant of SAF 205 (Mountain Hemlock). Transition between Pacific Silver Fir Series in Cascade-Sierran Montane Conifer Forest biotic community and Mountain Hemlock Series in Cascade-Sierran Subalpine Conifer Forest biotic community of Brown et al. (1998). Cascades-Cascade Crest Montane Forest Ecoregion, 4c (Pater et al., untitled; Thorson et al., 2003).
Historical Note and Mini-editorial: Forest vegetation presented in the two preceding slides (and to a large degree in the six preceding slides) made the case for the Gleasonian individualistic or continuum hypothesis of plant communities (Gleason, 1917, 1926). Gleason's individualistic concept of plant communities was anthetical to the Clementsian model of well-defined plant communities at multiple scales in space and time. An application of the continuum model was that of gradient analysis, "an ordination technique for the description of vegetation based on characteristics of the site rather than composition of the species" (Allaby, 1998). This technique can be most readily traced to early work in forests first by Gleason and later (and more influencial) by Whittaker (1956).
The gist (oversimplified for this purpose) of the gradient or continuum view as applied by workers like Whittaker (1956) was that resources and conditions for plants are so varied and the combinations of them so infinite (each resource and condition is arrayed along a continuum such that there are endless possible permutations) that any (each) plant community is unique (ie. one of a kind). The best that one could hope for is to combine key resources and growing conditions so as to able to describe habitat suitable for each or key species. The pictorial method devised to accomplish as the gradient diagram. Readers can find some of the classic examples of gradient diagrams in the standard plant ecology text by Barbour et al. (1999, ps. 582, 585, and 602). These have been most widely used for species in forest vegetation as for example when elevation is arrayed along one axis and moisture gradients along the other. The gradient diagram helps the student of vegetation visualize the environmental variables (eg. slope, aspect, soils, soil moisture, temperature, light, fire regime, elevation) and their combinations under which each plant species is capable of survival and reproduction.
Given the endless "things" that determine whether or not a species is fit for a given habitat (or whether any actual habitat will support a given species) it is unlikely that groups of plant species will routinely be found growing together. Again, most plant communities are unique and not repeatable (at least not greatly so). It follows that only individual communities can be described. There are not enough of the same (or even similar) plant communities to be able to identify or categorize plant communities into a group or "type". In other words, groups of species and their structure, age, arrangement, etc. (ie. plant communities) are not analogous to species. No Dr. Clements, groups of plants are not "superorganisms" or even "quasi-organisms"!
With the scientific likelihood that any given plant community is more-or-less unique and the relatively low probability of finding many others (perhaps none) like it, it would be of little use-- and maybe meaningless-- to attempt to "type" or otherwise categorize and descripe a given hypothetical plant community. This is the rationale of the continuum or individualistic hypothesis or concept ( see again, Barbour et al., 1999, ps. 23-24, 183-186) carried to its logical end.
This ecological logic argues: "If the continuum model is 'right on' then seeing, describing, mapping, etc. repeated units of vegetation (and at various spatial and temporal scales) is, at best, an abstraction (and an arbitrary one at that) and, at worst, an intellectual dilusion".
If the logical conclusion of the individualistic or continuum model is correct, it is a waste of endless time and effort (or nearly so) to try to lump, group, or whatever various units of vegetation into types (into any of the various types described elsewhere herein and in numerous texts and references). Indeed, if the individualistic paradigm is correct or closer to ecological truth than the discrete or organismic paradigm (Barbour et al., 1999, ps. 22-23, 182-183) it would seem that the Society of American Foresters (Eyre, 1980), Society for Range Management (Shiflet, 1994), and www.tarleton.edu/Departments/range were "lost causes". Why try to pen the wind?
The more relevant question, however, may be this: "Why do plant species form (development into) communities that are repeated in space and time and at several such units?". Why are there reognizable biomes and with numerous recgonizable units therein?
Then as an answer seems self-evident, some vegetation like that shown in the last few slides shows up. Maybe the questions on vegetation are as endless as the vegetation itself.
The Northern (sometimes, North) Cascades have been interpreted as a geological-physiographic province distinct from the High and Western Cascades south of the Columbia River and the adjoining Southern Washington Cascades (Franklin and Dryness, 1973, ps. 6,17-20). Matthews (1999, ps. 4, 6--8) followed this interpretative arrangement and pointed out that local topographic variation in the North Cascades is some of the most spectacular on Earth. Matthews (1999, p. 7-8) explained that the Cascade Crest and its rain shadow effects distinguishes East Slope from West Slope and produces fairly dramatic differences in natural vegetation within the province.
Presentations and descriptions of native vegetation of the North Cascades (and immediate contacts) in this portion began with lower elevation forests on the East Slope and progressed upward in elevation and into the "heart" of the Northern Cascades up over the Cascade Crest and down into low elevation forests of the West Slope. This constituted an interrupted photographic transect that followed the "ups and downs" of topography and elevation. Range vegetation thus "sampled" fell into three of the traditional life zones (Merriam, 1889): Transition (both Humid and Semiarid), Canadian and Hudsonian. Range types were either forests or subalpine mountain meadows, with many of the latter being invaded by trees. Consideration of range vegetation began with foothills forests near contact of the Okanogan Valley of the Columbia Plateau with the Northern Cascades. Physiographic unit is the Northern Cascade Mountains section of the Cascade-Sierra Mountains physiographic province (Feneman, 1931, ps. 422-430).
57. Northern Cascades Foothills Montane Forest- This north slope forest in the foothills of the Northern Cascades was an example of parklike, open-canopy ponderosa pine forest (or woodland) that by invasion by Douglas-fir was developing into a mixed western yellow pine-Douglas-fir closed canopy forest. All conifer regeneration was of Douglas-fir. At the same time numerous young pines (pole-size) were dying (some of which appeared to have been "shaded out" by Douglas-fir). Successful on-going invasion by Douglas-fir appeared to be largely due to absence of fire (due also perhaps to successionly recent changes in atmospheric phenomena (time scale of weather or recent climate). Increasing closure of forest canopy was serious threat to herbaceous understorey that was dominated by pinegrass (Calamagrostis rubescens) with local associate graminoid species varying from elk or Geyer's sedge (Carex geyeri), Thurber's needlegrass (Stipa thurberiana), Columbia brome (Bromus vulgaris), Cusick's bluegrass (Poa cusickii), Idaho fescue (Festuca idahoensis), and some Kentucky bluegrass (P. pratensis). The major forb overall was heartleafed arnica (Arnica cordifolia), but at on some local habitats arrowleaf balsamroot (Balsamorhiza sagittata) was dominant. Other forbs included some species of lupine (Lupinus sp.?), a vetch (Vicia sp.?), western yarrow (Achillea millefolium ssp. lanulosa), common dandelion (Taraxacum officinale), and various Lomatium species. The most common, though infrequent, shrub was common snowberry (Symphoricarpos albus).
Small pole-size Douglas-fir featured in foreground of these two photographs attested to recent invasion by Douglas-fir and development into a mixed (two-species) conifer forest that appeared to be further developing into a Douglas-fir forest. Ponderosa pine appeared to be on the way out, barring some intervening factor such as fire.
This SAF forest cover type could qualify as either Interior Ponderosa Pine (237) but going to Pacific Douglas-fir (229) due to unnatural fire cessation or, alternatively, as SAF 237, but as a seral type and going to SAF 229 which on this site would be a climax type (Eyre, 1980, ps. 91-92, 114-115).
This forest range plant community was in the Arid (actually, Semiarid) Transition (Life) Zone.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-17 (Pine-Douglas-fir Forest). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
58. Changing of the Guard- General structure and species composition (first photograph) and more details at closer distance (second photograph) of a north-slope, ponderosa pine-Douglas-fir-herbaceous understorey forest on which Douglas-fir was replacing--appeared to be excluding--ponerosa pine. Understorey was dominated by grasses including pinegrass (the dominant grass), Geyer's or elk sedge, Thurber's needlegrass, Columbia bromegrass, Cusick's bluegrass, Idaho fescue, along with some Kentucky bluegrass. Cheatgrass or downy brome was almost absent. The major forb--over all vegetation-- was heartleafed arnica, but at on some local habitats the dominant forb was arrowleaf balsamroot.
Note again the relative young age of Douglas-fir and fewer ponderosa pines as cohorts (ie. relatively lower regeneration of ponderosa pine). The apparent sapling in left foreground of second slide was actually two small trees: a Douglas-fir behind a ponderosa pine but with lower branches of the Douglas-fir in front of the ponderosa pine which otherwise was the only visible of these two conifer saplings. The accidental photographic illusion demonstrated the on-going regeneration of both conifer species though, again, Douglas-fir was reproducing at a faster rate and replacing ponderosa pine.
The most important distinguishing feature of this forest vegetation from the perspective of forest range was the continuous cover of herbage even where tree density was greatest. The herbaceous layer was present throughout this forest plant community even though species composition changed--often dramatically--from one local area to the next. Tree (canopy) cover was never so dense as to prevent development of a grazable understorey.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-17 (Pine-Douglas-fir Forest). SAF forest cover type could qualify as either Interior Ponderosa Pine (237) as climax vegetation and going to Pacific Douglas-fir (229) due to unnatural fire cessation or, alternatively, as SAF 237 as a seral type and going to SAF 229 which on this site would be a climax type (Eyre, 1980, ps. 91-92, 114-115). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
59. Foothill forest range- Two more views of a North Cascades foothills ponderosa pine-Douglas-fir forest with an herbaceous understorey dominated by pinegrass with elk of Geyer's sedge as associate. Other important grasses included Columbia brome, Thurber's needlegrass, Idaho fescue, Cusick's bluegrass, plus some naturalized Kentucky bluegrass. The dominant forb (conspicuous in the second of these two photographs) was arrowleaf balsamroot. Heartleafed arnica was the major forb averaged across the range vegetation, but there were local habitats where some unidenfied species of lupine (Lupinus sp.) was the dominant or associate forb.
Structure of this forest was typical in having local openings (smaller areas devoid of trees; "natural clearings"). Untypical (and unnatural), however, was the invasion of many of these small forest openings by Douglas-fir. The forest opening in the second of these two slides was not--for the time being--being invaded by Douglas-fir. Forest openings were considered further in the next set of photographs.
This mesophytic form or phase of foothill forest was on a predominately north-facing slope.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-17 (Pine-Douglas-fir Forest). SAF forest cover type could qualify as either Interior Ponderosa Pine (237) as climax vegetation and going to Pacific Douglas-fir (229) due to unnatural fire cessation or, alternatively, as SAF 237 as a seral type and going to SAF 229 which on this site would be a climax type (Eyre, 1980, ps. 91-92, 114-115). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
60. Openings in a once-open forest range- Natural forest openings are areas in forests that for whatever reason(s) are devoid of trees and without canopy cover from overhanging branches of adjacent trees. Such forest openings were part of the structure of the ponderosa pine-Douglas-fir-herbaceous understorey forest that was used as a representative example of forest range in the foothills of the North Cascades. Several such natural openings were present (and featured) in this description of range vegetation.
Althouth herbaceous plants, especially grasses and sedges, dominated forest openings (and the lower forest layers in general) one shrub that was present at quite limited cover and occurrence was common snowberry. An example was shown in the first of these two slides (at right margin of foreground). It was revealing of the extent of Douglas-fir invasion in this range vegetation that a large seedling or small sapling had emerged (and was visible) sfrom the center of the snowberry. The second photograph featured the edge of a natural opening in which a Lupinus species was the locally dominant forb.
It was remarked previously, and bears reemphasizing, that the herbaceous layer(s) of vegetation was present consistently throughout this forest range.For example, the grassy nature of the lower layer was conspicuous under fairly dense stocking of Douglas-fir in the background of the first slide.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-17 (Pine-Douglas-fir Forest). SAF forest cover type could qualify as either Interior Ponderosa Pine (237) as climax vegetation and going to Pacific Douglas-fir (229) due to unnatural fire cessation or, alternatively, as SAF 237 as a seral type and going to SAF 229 which on this site would be a climax type (Eyre, 1980, ps. 91-92, 114-115). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
61. Grass up to the bark- Photo-plot of interior of ponderosa pine-Douglas-fir-"grassy" understorey forest on a predominately north-facing slope in foothills of Northern Cascades. Young trees (including trunk in foreground) were mostly Douglas-fir. Many of the branches of these trees (again, foremost foreground tree) supported abundant growth of wolf lichen (Letharia vulpina). This "plot" featured the characteristic (and continuous) herbaceous layer dominated by pinegrass grass and elk sedge. Other major grasses included Thurber's needlegrass, Columbia bromegrass, Cusick's bluegrass, Idaho fescue, and some naturalized Kentucky bluegrass. Cheatgrass was the same as absent as was bluebunch wheatgrass. The prominent forb in this photograph was western yarrow. Viewers attention was drawn to the present of herbaceous plants under conifers, usually next to tree boles ("up to the bark" as it were).
The only shrub of any consequence was common snowberry with a representive at right margin of foreground.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-17 (Pine-Douglas-fir Forest). SAF forest cover type could qualify as either Interior Ponderosa Pine (237) as climax vegetation and going to Pacific Douglas-fir (229) due to unnatural fire cessation or, alternatively, as SAF 237 as a seral type and going to SAF 229 which on this site would be a climax type (Eyre, 1980, ps. 91-92, 114-115). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
62. Foliage galore- Interior of a Northern Cascades foothills ponderosa pine-Douglas-fir-herbaceous understorey forest showing the amazing diversity of plant life in this forest range. Boughs of Douglas-fir, the dominant (at least becoming so) conifer, were just a short distance above a species-rich sward dominated by pinegrass and with Geyer's or elk sedge, Columbia bromegrass, Thurber's needlegrass, Cusick's bluegrass, Idaho fescue, and some naturalized Kentucky bluegrass. Forbs included heartleafed arnica, the most common species forest-wide, a species of lupine, and, sticking out like so many diamonds in as many goat's anuses, naturalized common dandelion (Taraxacum officinale).
Encroachment of the invading Douglas-fir into natural forest openings was obvious and constituted an eventual threat to the herbaceous layer(s) of the species- and range feed-rich north-slope forest. Cessation of fire was permitting what was probably unnaturally (abnormally) greater cover of Douglas-fir in a forest range cover type that is regarded by many authorities as a fire type. Increasing canopy cover of Douglas-fir also threatens ponderosa pine which is probably the natural co-dominant conifer for this forest cover type. This threat to integrity of natural forest range vegetation was dealt with further below.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-17 (Pine-Douglas-fir Forest). SAF forest cover type could qualify as either Interior Ponderosa Pine (237) as climax vegetation and going to Pacific Douglas-fir (229) due to unnatural fire cessation or, alternatively, as SAF 237 as a seral type and going to SAF 229 which on this site would be a climax type (Eyre, 1980, ps. 91-92, 114-115). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
63. Plenty of feed in this understorey (or pinegrass under the pines-- and Douglas-firs)- The herbaceous layer(s) of a Northern Cascades foothill (north slope) ponderosa pine-Douglas-fir-"grassy" understorey forest. This grassland-like sward was composed of numerous grasses including pinegrass, the overall herbaceous dominant, Thurber's needlegrass, Columbia brome, Idaho fescue, Cusick'sbluegrass, and, locally abundant, naturalized Kentucky bluegrass along with Geyer's or elk sedge which was frequently the associate herbaceous species. The most common forb on a forest-wide basis was heartleafed arnica some leaves of which were featured in the foreground of the first of these two photographs.
Needles of both conifer species were in the sward layer(s) of this foothill forest range. Pine straw was especially noticable in the second of these two photo-quadrants. Also obvious in the second photograph was movement of grass inflorescences by wind (visible as blurred images of pinegrass and Thurber's needlegrass panicles).
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-17 (Pine-Douglas-fir Forest). SAF forest cover type could qualify as either Interior Ponderosa Pine (237) as climax vegetation and going to Pacific Douglas-fir (229) due to unnatural fire cessation or, alternatively, as SAF 237 as a seral type and going to SAF 229 which on this site would be a climax type (Eyre, 1980, ps. 91-92, 114-115). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
64. "A clear and present danger" (or is it?)- Invasion and encroachment of Douglas-fir seedlings into natural openings in a Northern Cascades foothill (north slope) ponderosa pine-Douglas-fir-"grassy" understorey forest range. Herbaceous layer(s) were generally dominated by pinegrass with elk or Geyer's sedge as associate herb. Other important grasses included Columbia brome, Thurber's needlegrass, Cusick's bluegrass, Idaho fescue, naturalized Kentucky bluegrass and, less commonly, cheatgrass. Locally common to dominant forbs included heartleaf arnica, arrowleaf balsamroot, and lupine. Common snowberry was occasionally present and constituted a sporatic shrub layer in this forest range vegetation.
Abnormally high levels of regeneration of Douglas-fir was attrributed--at least partially--to absence of fire from what is to some degree a fire type. Absence of fire and, almost assuredly, other factors (perhaps including limited elk grazing) was permitting Douglas-fir reproduction to increase to excessive levels of canopy cover and into heretofore natural forest openings that supported botanically diverse local herbaceous stands and rich feed sources for range ungulates, including cattle, mule deer, and elk.
The precise term of invasion was employed throughout much of this discussion in an equally precise usage. Invasion: "Invasion is the movement of one or more plants from one area into another and their establishment in the later It is thus the complete and complex process of which migration, ecesis , and competition are the essential parts. It is going on at all times and in all directions" (Weaver and Clements, 1938, p. 166). Complete invasion refers to the condition "when the movement of invaders into a community is so great the the original occupants are driven out" (Weaver and Clements, 1938, p. 168). Presented in these two photo-plots (and several others above) is the ecological condition in which Douglas-fir (abnormally high cover thereof) is in early stages of threatening complete invasion of this ponderosa pine-Douglas-fir-herbaceous species forest with eventual replacement of much of the more natural cover of herbaceous species.
This unnatural condition is an "in the works" potential threat to ecological integrity of this forest range ecosystem with the likely loss of much of the range feed resource and plant species diversity. The "delicate balance" of a natural ecosystem "hangs in the balance". Solution: bring out the drip touches. In western spirit, "Powder River let her buck!" Or in the parlance of the 1960s draft card-burning protesters, "Burn baby, burn!" And it better be sooner rather than latter: burn now or pay later.
Another interpretation: Daubenmire and Daubenmire (1968, ps. 23-25) recognized a climax Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) that had included "no coniferous trees other than the Pseudotsuga" and with "an udergrowth that is an essentially shrub-free sward dominated outstandingly by Calamagrostis rubescens". Daubenmire and Daubenmire (1968, p. 25) cited other workers who described a "Conifer-Calamagrostis only" type which with "extensive and repeated burning" eliminated Douglas-fir seed sources thereby allowing ponderosa pine to retain dominance with pinegrass. This latter forest vegetation was interpreted as a ponderosa pine-pinegrass cover type within the Pseudotsuga menziesii-Calamagrostis reubescens habitat type. Daubenmire and Daubenmire (1968, p. 25) reported this habitat type in the "eastern foothills of the Cascade Mountains in Washington". From perspective of Daubenmire and Daubenmire (1968) the forest range plant community presented here was climax vegetation and ponderosa pine was a seral species that was gradually being replaced by the climax Douglas-fir. Thus, the invasion shown and described above was natural and normal, representing plant succession. This begs the question as to how natural or unnatural and fire or absence of fire were in this forest type. The frequent occurrence of common snowberry was also a ecological "fly in the oitment" of the "essentially shrub-free sward". Snowberry was common on the specific range on which this ponderosa pine-Douglas-fir-"grassy" layer forest occurred. In fact, there was an adjoining range plant community of the climax Pinus ponderosa-Symphoricarpos albus habitat type (Daubenmire and Daubenmire, 1968, ps. 8-12).
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-17 (Pine-Douglas-fir Forest). SAF forest cover type could qualify as either Interior Ponderosa Pine (237) as climax vegetation and going to Pacific Douglas-fir (229) due to unnatural fire cessation or, alternatively, as SAF 237 as a seral type and going to SAF 229 which on this site would be a climax type (Eyre, 1980, ps. 91-92, 114-115). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
65. The herbaceous dominant and a local associate- Pinegrass and an unidentified lupine (Lupinus sp.?) in the herbaceous understorey (one or more layers) of a north slope, Northern Cascades foothills ponderosa pine-Douglas-fir forest range.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer). Phenological stages: full-bloom (anthesis) in pinegrass, pre-bloom in lupine.
66. Sexual reproduction in the herbaceous dominant- Panicle of pinegrass in the herbaceous layer of a ponderosa pine-Douglas-fir-herbaceous understorey forest range in foothills of Northern Cascades. There is some variation in panicle branching among individuals of pinegrass. Some panicles are more fully expanded by having primary branches coming off of the central axis at wider angles (see next two photographs) whereas in other panicles (the one shown here) primary branches remain closer to the central axis.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer). Anthesis.
67. Panicles of pinegrass- Morphology and details of the inflorescence of Calamagrostis rubescens.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer). Anthesis stage of phenology.
68. Poor representation, but present nonetheless- One--the best one--of only a few panicles (an early blooming specimen) of Columbia bromegrass in the herbaceous layer(s) of a north slope, Northern Cascades foothills ponderosa pine-Douglas-fir forest range. On this range this species was never even an associate species, but it was consistently present at low densities and cover in the sward of this range plant community.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer).
69. Thurber's needlegrass (Stip thurberiana)- Some of a very few examples of Thurber's needlegrass that were in bloom on a north slope, Northern Cascades ponderosa pine-Douglas-fir-herbaceous layers forest range. This grass rated the ranking of associate species on local habitats, but it was clearly second to elk or Geyer's sedge as an associate. Fortunately for this lesson, however, it was more sexually reproductive than the sedge which failed to make its full contribution to the lesson.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer). Phenological stage presented here was grain-ripe, pre-shatter stage.
70. Like as to a plant- Population of wolf lichen (Letharia vulpina) on a limb of Douglas-fir in the Northern Cascades ponderosa pine-Douglas-fir-herbaceous understorey foothill forest range described above. Lichen refers to an organism that is a symbiotic combination of algae and fungus existing in mutualism (mutual dependence) and so biologically integrated that this organism becomes unique from its component organisms and is interpreted as a separate species (even though it is two species living together).
What to call, regard, or interpret lichen is a more basic--though no less confusing--problem. In the "good ole days" of Botany and Zology when every organism was either plant or animal, lichen clearly was a plant. With the newer five kingdom (and probably more on the way) system of classification things are not so simple. In their textbook entitled Biology of Plants Raven et al. (1992) avoided defining plant. Raven et al. (1992, p. 11) seemingly managed to justify inclusion of organisms that are not plants in Biology of Plants with this rationalization: "Although we do not regard algae, fungi, bacteria, or viruses as plants, and shall not refer to them as plants in this book, they are included here because of tradition, and because they are normally considered as part of the botanical portion of the curriculum..." . Whatever lichens are, they were part of the range vegetation this example of Northern Cascades foothill forest and were "included here because of tradition..."
Wolf lichen was also included with treatment of Rocky Mountain forest range (Northern Rocky Mountains) where this not-referred-to-as-a-plant range plant was covered in more detail. This species was not discussed further in this chapter.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect of vegetation.
71. Pristine combination- An example of a ponderosa pine-common snowberry foothill forest range. This natural vegetation developed conterminous with the ponderosa pine-Douglas-fir-"grassy" layer foothill forest presented immediately above. Both of these examples were on the same range (same grazing unit; no fences separated them) with the only differences being range sites and habitat types. This was an example of the climax Pinus ponderosa-Symphoricarpos albus habitat type (association) of Daubenmire and Daubernmire (1968, ps. 8-12) and recognized as an association by Franklin and Dryness (1973, ps. 173). Herbaceous species were limited to perimeters of snowberry plants which otherwise effectively excluded other plants. The major herbaceous range plants were blue wildrye (Elymus glaucus) and various Lomatium species. Pinegrass, Geyer's or elk sedge, and Thurber's needlegrass were less common than blue wildrye.
The Society for Range Management rangeland cover type description (Shiflet, 1994) interpreted this as the most mesic climax form of Ponderosa Pine Shrubland (109).
This forest community was primarily browse range with herbaceous range plants clearly minor compared to snowberry. This vegetation was contiguous with the forest community presented next (immediately below) as well as the ponderosa pine-Douglas -fir-herbaceous layers forest described above.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-10 (Western Ponderosa Forest). SAF 237 (Interior Ponderosa Pine). SRM 109 (Ponderosa Pine Shrubland). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
72. Another climax combination- Ponderosa pine-mallow ninebark foothill range. This relatively limited example of a less common range type was contiguous with the ponderosa pine-Douglas -fir-herbaceous layers forest and the ponderosa pine-common snowberry forest described above. The forest range vegetation featured now was an example of the climax Pinus ponderosa-Physocarpus malvaceus habitat type (association) of Daubenmire and Daubernmire (1968, ps. 12-13) and recognized/cited by Franklin and Dryness (1973, ps. 173). Blue wildrye and Lomatium species were the major range grass and range forb. There was also some pinegrass, Thurber's needlegrass, Columbia bromegrass and elk sedge among graminoids. The second most abundant forb was heartleafed arnica.
Like the preceding ponderosa pine-common snowberry the ponderosa pine-mallow ninebark type is primarily browse range.
Washington State University's Colockum Unit, Chelan County, Washington. June, estival aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-10 (Western Ponderosa Forest). SAF 237 (Interior Ponderosa Pine). SRM 109 (Ponderosa Pine Shrubland). Yellow Pine Series (122.55) in Cascade-Sierran Montane Conifer Forest biotic community (122.5) of Brown et al. (1998, p. 37). Columbia Plateau- Okanogan Valley Ecoregion, 10m (Environmental Protection Agency, undated).
07a 320
73. Common snowberry (Symphoricarpos albus)- Plant of common snowberry, the dominant shrub and single-species component of a shrub layer, in a ponderosa pine-snowberry forest range in foothills of Northern Cascades.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer). Full-bloom stage of phenology.
74. Flowering leader of common snowberry- A sexual shoot on the plant introduced in the preceding slide. Ponderosa pine-snowberry forest range in foothills of Northern Cascades.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer). Full-bloom phenological stage.
75. Snowberry flowers- Details of inflorescences of common snowberry. These were on the plant presented above that was in shrub understorey of a ponderosa pine-snowberry browse range in foothills of Northern Cascade.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer).
75. Western virgin's bower (Clematis ligusticifolia)- Another of the showy shrubs in the foothills of the Northern Cascades is this Clematis species which prefers mesic habitats such as part of riparian range vegetation, on north and east slopes, and in protected coves under ponderosa pine like the examples presented above.
Washington State University's Colockum Unit, Chelan County, Washington. June (early summer).
The montane and subalpine (vs. foothills) forests within the Northern Cascades treated below were organized like an interrupted photo-transect that was recorded and described periodically along "ups and downs" of elevation and on specific range environments (eg. range and forest sites, habitat types, cover types and variants). These involved the three major zones of climax forest vegetation of Franklin and Drynness, 1973, ps.70-108 passim): 1) Western Hemlock (Tsuga hetrophylla) Zone, 2) Pacific Silver Fir or (Abies amabilis) Zone and 3) Mountain Hemlock (Tsuga mertensiana) Zone. The Tsuga heterophylla Zone included subclimax Douglas-fir (Pseudotsuga menziesii) foressts which began this photographic survey of range vegetation in the North Cascades above the foothills.
Douglas-fir, including both Pacific or coast and interior, Rocky Mountain or blue varieties (Pseudotsuga menziesii var. menziesii and P. menziesii var. glauca, respectively), has historically been regarded as a seral or subclimax forest species that colonizes or pioneers disturbed sites especially where mineral soil is exposed (fire, flood, blowdown, etc.). Douglas-fir is such a long-lived tree that individuals persist from initial or early seral stages into the climax forest vegetation. As a consequence of the silvics or life cycle of Douglas-fir forests typically have scattered old--thus relatively old--trees of this species. Douglas-fir is the single most important lumber species in the United States of America and is typically managed silviculturally for what it is: a marvelous lumber crop. Examples of such forest management were presented variously throughout this publication. Forests in which Douglas-fir comprises most of the stocking are subclimax, but these include several major forest cover types which were also presented periodically.
Although Douglas-fir (and forest vegetation dominated by it) is subclimax such range vegetation is so persistent and economically important that it merits coverage as an important forest range type. Furthermore, it remains a matter of debate as to whether Douglas-fir-dominated (or defined) forests that developed in stream drainages--as riparian vegetation (or part of it) or forest just beyond the riparian zone riparian--is in successional fact seral, subclimax, climax, postclimax, (or whatever).
Coverage of range vegetation within the montane and subalpine zones of the Northern Cascades Mountains began with Douglas-fir forests that developed within streams drainages just beyond the wet riparian zone. The valley of Early Winters Creek and the forest range vegetation that developed in it served as the example of this forest plant community. This forest range type was in the lower part of the Canadian Life Zone (Merriam, 1889).
76. Forming a forest- On an easterly slope a Pacific Douglas-fir forest (Pseudotsuga menziesii var. menziesii) was replacing an open, herbaceous nderstorey ponderosa pine forest or woodland.as the "new"forest". Perhaps Douglas-fir-dominated forest is the climatic climax for this forest site whereas the existing open-canopy ponderosa pine forest was either seral to Douglas-fir or a fire type (= pyric climax). Fire is, however, one of the main (if not the main) destructive agent that facilitates establishment of Douglas-fir by creating mineral soil surface conditions conducive to germination, emergence, and establishment of Douglas-fir seedlings (Eyre, 1980, p. 106). Franklin and Dryness (1973 ps.) interpreted most Douglas-fir-dominated forests in the Cascade Ranges as seral (but with some individuals of this long-lived conifer persisting into climax to be a component thereof) as part of the climax Tsuga heterophylla zone. The Pacific Douglas-fir forest cover type (SAF 229) is sometimes climax on drier sites and under certain topographic conditions (Eyre, 1980, p. 107).
Douglas-fir was forming a "pure" single-conifer species on this site, but a shrub understorey had persisted dominated by grouseberry or whortleberry (Vaccinium scoparium) and thinleafed blueberry, tall bilberry, or big huckleberry (V.membranaceum) along with sticky laurel, of tobaccobush (Ceanothus velutinus). Herbaceous species were limited, but grasses were of Poa and Bromus species, none of which were blooming and identifiable to this "alien" photographer.
Okanogan National Forest (above Early Winters Creek), Okanogan County, Washington. June (early summer), estival aspect. FRES No. 20 (Douglas-fir Forest Eciosystem). K-11 (Douglas-fir Forest). SAF 229 (Pacific Douglas-fir). No unit in Brown et al (1998, p.37): should be a Douglas-fir Series in Cascade-Sierran Montane Conifer Forest biotic community (122.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
77. Beneath mammy and pappy- Douglas-fir regeneration in the shade of adult Douglas-fir is typically uncommon in this species (Eyre, 1980, ps. 106-107) that was generally rated as Intermediate in tolerance (Wenger, 1984,, p. 30). Nonetheless, there was reproduction of the next generation of Douglas-fir on this lower east slope in valley of Early Winters Creek in the lower North Cascades. Species in the lower layers of this forest vegetation included grouseberry or whortleberry and thinleafed blueberry, tall bilberry, or big huckleberry as major shrubs with perennial grasses being mostly bluegrasses and bromes.
Okanogan National Forest (along Early Winters Creek), Okanogan County, Washington. June (early summer), estival aspect. FRES No. 20 (Douglas-fir Forest Eciosystem). K-11 (Douglas-fir Forest). SAF 229 (Pacific Douglas-fir). No unit in Brown et al (1998, p.37): should be a Douglas-fir Series in Cascade-Sierran Montane Conifer Forest biotic community (122.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
78. Layers in the woods- A composite photograph that showed species composition and structure of a creek bottom Douglas-fir-dominated forest with regeneration of Douglas-fir comprising part of an upper woody far below the canopy. A second (lower) shrub layer was made up of of mainly Ericaceous species, especially grouseberry or whortleberry and thinleafed blueberry, tall bilberry, or big huckleberry. Herbaceous plants were uncommon and consisted mostly of Poa, Bromus,and Carex species, none of which this "stranger to these parts" photographer could identify from vegetative features.
Okanogan National Forest (along Early Winters Creek), Okanogan County, Washington. June (early summer), estival aspect. FRES No. 20 (Douglas-fir Forest Eciosystem). K-11 (Douglas-fir Forest). SAF 229 (Pacific Douglas-fir). No unit in Brown et al (1998, p.37): should be a Douglas-fir Series in Cascade-Sierran Montane Conifer Forest biotic community (122.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
79. Ground level- Woody understorey of a creek bottom Douglas-fir forest. This particular view of the lower levels of this forest range lacked the layer dominated by saplings of Douglas-fir and some of the other shrub species present in the lower shrub layer as was show in preceding slides.
Okanogan National Forest (along Early Winters Creek), Okanogan County, Washington. June (early summer), estival aspect. FRES No. 20 (Douglas-fir Forest Eciosystem). K-11 (Douglas-fir Forest). SAF 229 (Pacific Douglas-fir). No unit in Brown et al (1998, p.37): should be a Douglas-fir Series in Cascade-Sierran Montane Conifer Forest biotic community (122.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
80. One for the textbooks- Douglas-fir forest range with reproduction of Douglas-fir and an herbaceous layer comprised of various grasses including pinegrass, blue wildrye, along with Bromus and Poa that were not blooming and which this "first-time visitor" author could not identify vegetatively. There was also some Carex species present. In other parts of the herbaceous layer(s) forbs were dominant, often exclusive, species. An example of this form of Douglas-fir-herbaceous species forest had developed conterminous with the forest vegetation shown here (just to the far right of the forest plant community shown in this photograph). This out-of-view Douglas-fir-forb forest range was presented in slides immediately below.
The shrub in left foreground was a small common snowberry. Shrubs were typically wanting from the "grassy" layer form of Douglas-fir forest represented here.
An example of the "grassy" understorey form of Douglas-fir forest range. This forest vegetation was similar to the Pseudotsuga menziesii-Calamagrostis reubescens habitat type (association) of Daubenmire and Daubenmire (1968, ps. 23-25) that was described above.
A doe mule deer was standing just out of the shade in the background (right of center) as if to underscore importance of this forest vegetation as valuable game range.
Okanogan National Forest (above Early Winters Creek), Okanogan County, Washington. June (early summer), estival aspect. FRES No. 20 (Douglas-fir Forest Eciosystem). K-11 (Douglas-fir Forest). SAF 229 (Pacific Douglas-fir). No unit in Brown et al (1998, p.37): should be a Douglas-fir Series in Cascade-Sierran Montane Conifer Forest biotic community (122.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
81.Different form of herbaceous layer under Douglas-fir- Douglas-fir forest range with an herbaceous layer dominated by the composite forb, palmateleaf colt'sfoot (Petasites frigidus var. palmatus). This "sample" of Douglas-fir forest vegetation was immediately adjacent to the Douglas-fir-"grassy" understorey described in the preceding slide.
This was an uneven-aged stand of Douglas-fir indicating relatively recent regeneration. This was consistent with Douglas-fir reproduction in the "grassy" portion of this same forest as shown in the photograph immediately above.
Okanogan National Forest (above Early Winters Creek), Okanogan County, Washington. June (early summer), estival aspect. FRES No. 20 (Douglas-fir Forest Eciosystem). K-11 (Douglas-fir Forest). SAF 229 (Pacific Douglas-fir). No unit in Brown et al (1998, p.37): should be a Douglas-fir Series in Cascade-Sierran Montane Conifer Forest biotic community (122.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
82. Colony of colt'sfoot- Part of the herbaceous layer of the Douglas-fir-herb species forest range introdued in the two preceding slides was dominated by palmateleaf colt'sfoot. Hitchcock and dronquist (1973, p. 541) recognized two varieties of Petasites frigidus. This is the palmateleaf variety: Petasites frigidus var. palmatus. Colt'sfoot is in the groundsel tribe (Senecioeae) of the Compositae. The commonness--often dominance--of the composites in forest and rangeland vegetation was again evident in the native plant community described here.
Okanogan National Forest (above Early Winters Creek), Okanogan County, Washington. June (early summer). Immediate post-bloom to early fruit phenological stage.
83. Green feet in the forest- Leaves of palmateleaf colt'sfoot in the colony presented in the preceding photograph.Okanogan National Forest (above Early Winters Creek), Okanogan County, Washington. June (early summer).
84. Blooming colt's foot- Detail of sexual shoot and infloresences of the colt'sfoot colony in understorey of Pacific Douglas-fir forest described above.
Okanogan National Forest (above Early Winters Creek), Okanogan County, Washington. June (early summer). Immediate post-bloom to early fruit phenological stage.
85.A prize in the colt herd- An individual of the orchid, western ladies tresses (Spiranthes porrifolia= S. romanzoffiana var. porrifolia) inside the colony of coltsfoot shown above. The two leaves behind the orchid were of a species of Arnica (perhaps A. cordifolia) which is, like colt'sfoot, in the groundsel tribe of Compositae. Appropriately enought the largest family (Orchidaceae) and second largest family (Compositae) of flowering plants (angiosperms) were represented in this local herbaceous layer of the Douglas-fir-dominated forest treated at this point.
While there are relatively few Spiranthes species in the greater Pacific Northwest region this is a widely distributed genus of forb with other Spiranthes species native to the prairies to the east. Two other species of ladies tresses were presented in the chapter on Tallgrass Prairie (Interior).
Okanogan National Forest (above Early Winters Creek), Okanogan County, Washington. June (early summer). Peak bloom stage.
86. Trunks of the dominant- Two individual trunks of mature Pacific Douglas-fir (Pseudotsuga menziesii var. menziesii ) growing in the creek bottom Douglas-fir-dominated forest vegetation presented above (along Early Winters Creek). Even with early to mid-morning light coming directly into this forest there was still shade. Shade (mostly from Douglas-fir boughs) precluded showing bark on the boles to full advantage, but it was the best possible under the prevailing conditions-- and it illustrated importance of light (or, the opposite, shade) in forests dominated by Douglas-fir.
Incidentially, even with this degree of shade (and this was as sunny as this locale gets) there were low shrubs (mostly Ericaceous) in lower levels of this forest. This documented importance of this natural vegetation as range (on this range, mostly mule deer)..
Okanogan National Forest (above Early Winters Creek), Okanogan County, Washington. June (early summer).
Subalpine fire western red cedar, and western hemlock are Very Tolerant Engelmann spruce, Mountain hemlock and Pacific silver fir are Tolerantis Douglas-fir an western white pine are intermediate in tolerance (Wenger, 1984, p. 6).
"Uphill" (higher in elevation) and farther from a stream, its relief, and its influences (drier, shallower soil, greater exposure to light, etc.) in the Northern Cascades Douglas-fir remains an important, but not longer a dominant, component of forest plant communities. The long-lived Douglas-fir persists in later (more advanced) successional stages perhaps as a member of the climax forest. On such forests in the Northern Cascades Douglas-fir is frequently a member of climax forest vegetation dominated or becoming dominated by such conifers as Engelmann spruce, Pacific silver or Cascades fir, subalpine fir (Abies lasiocarpa), mountain hemlock or various combinations of these species.
An example of this forest vegetation was presented next. This forested range community was in the upper reaches of the Canadian Life Zone (Merriam, 1889). It was the East Slope of North Cascades meaning east of the true Cascade Crest (Matthews, 1999, ps.7-8). It was in the Abies amabilis zone of Franklin and Dryness (1973, ps.93), meaning that Pacific silver or Cascade fir is the zonal (=climatic climax) dominant conifer.
87. Douglas-fir-Engelmann spruce-Pacific silver fir montane forest- Outer view of a subclimax Engelmann spruce-Pacific silver (with persistent seral Douglas-fir) forest on a southerly slope. This was a cool temperate forest well below subalpine environments. Douglas-fir (eg. largest tree in this view) persisted into this forest from previous sucessional stages, but there was no regeneration of this conifer. Instead most regeneration was Pacific silver fir with some reproduction of Engelmann spruce. The dominant shrub in the understorey was tobacco-brush. Sitka mountain-ash (Sorbus sitchensis var. sitchensis) and bearberry manzanita or kinnikinnick were associate shrubs. The most abundant herbaceous species was broadleaf or big leaf lupine (Lupinus latifolius). Grasses and grasslike plants were limited to openings.
This was as close as the photographer could get to a view of physiogonomy of this forest plant community in a generally continuous forest cover.
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer), early estival aspect. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Foorest biotic community (!22.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
88. Shrubs and a "die-hard" Douglas-fir- Closer-in view of the forest range vegetation presented in the preceding slide. This view of the lower trunk of the old Douglas-fir showed to better advantage tobacco-brush, the largest and tallest shrub (outer foreground). The shrubs behind tobacco-brush were plants of bearberry manzanita or kinnikinnick. The partial crown of a conifer sapling along margin of left foreground was Pacific silver fir.
Okanogan National Forest, Okanogan County, Washington. June (early summer), early estival aspect. The various units of vegetation classification were not presented here because this was such a restricted view (see captions immediately above and below)
89. Forest edge- Outer perimeter of a south slope Douglas-fir-Engelmann spruce-Pacific silver fir forest in Northern Cascades. Edges of plant communities (or, for that matter, edges of single-species fields of crops) are different from interiors of these same plant communities (populations of field or orchard crops). Edges where two (or more) different communities or populations meet are some of the most interesting assemblages of plants, and typically the most productive for ecotonal species like certain species of wildlife. Shown here was the outermost edge of a later seral stage, perhaps subclimax, forest of Engelmann spruce-Pacific silver fir with Douglas-fir persisting since its establishment in an earlier, maybe pioneer, stage of plant succession.
This view "sets the stage" for a Vegetation Science visual laboratory farther into this forest range plant community as shown in the photographs that follow. The range vegetation on this perimeter of the forest consisted of shrubs, forbs, grasses,and sedges. Shrubs were featured in this photograph with other forest range plants dealt with in succeeding slides. Dominant (major) shrubs were tobacco-brush (tobaccobush) or sticky laurel and bearberry manzanita or kinnikinnick (Arctostaphylos uva-ursi).
The root wad of the wind-toppled tree in midground delineated a recently created forest gap above while the small crater served as a "tree cradle" which in this second or third growing season post- windthrow was filled with cheatgrass or downy brome.The conspicuous pale- to gray-green grass around perimeter of the crater was intermediate wheatgrass (Agropyron intermedium), a Eurasian perennial introduced as a naturalized range forage crop.
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer), early estival aspect. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Foorest biotic community (!22.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
90. Species composition and structure of a Douglas-fir-Engelmann spruce-Pacific silver fir forest- Attempted "all-in-one" view of the subclimax cool temperate forest introduced above. The different layers of forest range vegetation were fairly well-defined in this perspective. Largest tree (left foreground) was the old Douglas-fir shown above with tobacco-brush, bearberry manzanita, and Sitka mountain-ash as major shrubs, but at far right plants of pinemat manzanita (Arctostaphylos nevadensis) "joined the staff". Herbaceous species were featured farther below. Most of the larger trees in center to right foreground were Engelmann spruce. There was some reproduction of this conifer, but most regeneration was of Pacific silver fir. Both of these species were rated as Tolerant (Wenger, 1984, p. 6). There was essentially no regeneration of Douglas-fir which is generally rated as Intermediate in tolerance (Wenger, 1984, p. 6).
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer), early estival aspect. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Foorest biotic community (!22.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
91. First and second on the scene meet the last (in both senses) to arrive- Early seral--probably, pioneer--Douglas-fir and cimax or, maybe, subclimax Engelmann spruce stand grandly by as climax scrubby Pacific silver or Cascades fir slowly starts to take control of this cool temperate forest on East Slope of the Norther Cascades. In the first of these two photographs Douglas-fir was represented by the three larger trunks in the left foreground to midground while Engelmann spruce was represented by the three larger trunks in center to right foreground. The second of these two photographs took up with the three Engelmann spruce and featured to the right of these a large Douglas-fir and an Engelmann Spruce behind it.
Although the ground was littered with cones of Engelmann spruce most regeneration was that of Pacific silver fir.This included large seedlings such as the cute little one in front of the largest Engelmann spruce (center foreground of first slide; left margin of foreground in second slide) to saplings with their characteristic horizontal limbs often with an upward sweep.
Shrubs shown in the first photograph included tobacco-brush, bearberry manzanita or kinnikinnick, and pinemat manzanita. Grasses and sedges were in early stages of growth and this author could not identify them from vegetative characteristics other than to determine some as Poa, Bromus, and Carex species.
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer), early estival aspect. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Foorest biotic community (!22.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
92. Lower layers- Ground (lower herb and shrub) and intermediate layers in a Douglas-fir-Engelmann spruce-Pacific silver fir col temperate forest on East Slope of Northern Cascades. Upper surface of soil included conifer needles and abundant cones of Engelmann spruce. The green clumps included Carex, Bromus, and Poa species which this interloper phototgrapher could not identify to species in their pre-bloom stages. Conifer seedlings and saplings were almost exclusively Pacific silver fir. Neither Douglas-fir nor Engelmann spruce had noticable regeneration. This was the typical pattern in progression of conifer species with advancing succession as reported by Johnson and O'Neil (2001, p. 31). Large trunks of trees in the background were both Engelmann spruce (mostly) and Douglas-fir (largest tree at right).
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer), early estival aspect. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Foorest biotic community (!22.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
93. Mat to the butts (almost)- Another "photo-plot" of the lower layer(s) of the Douglas-fir-Engelmann spruce-Pacific silver fir forest (East Slope of Northern Cascades) being described here. Pinemat manzanita formed a local low, matlike layer of vegetation as part (and over part) of the understorey of this late seral or, perhap smore precisely, sublcimax forest. This layer grew up to the butts of Douglas-fir trunks. The background was composed mostly of invading Pacific silver fir, the climax dominant of this forest site and the only conifer that was regenerating (at least to any extent).
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer), early estival aspect. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Foorest biotic community (!22.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
94. The climax comes in- Exterior view of a Douglas-fir-Engelmann spruce-Pacific silver fir cool temperate forest showing invasion by Pacific silver or Cascades fir, the climax dominant conifer for this forest site. Almost all regeneration was of Pacific silver fir. The largest trees were Douglas-fir followed closely by Engelmann spruce. Douglas-fir is an early seral--perhaps pioneer conifer--that lives to such old age (numerous individuals live to be 300-500 years old) that individuals persist into the vegetation of climax forests. Douglas-fir reaches immense sizes (both height and diameter) due to its old age and rapid growth so that trees of this species frequently "over-shadow" almost all others in forests of the Pacific Northwest. Engelmann spruce is not all that far behind Douglas-fir in age and dimension, but it is appears later in the forest sere and typically persist as a component of the climax forest as well. Pacific silver or Cascades fir is the true climax conifer that arrives latest on the "seral scene" of many forest sites, especially those just below the prime elevational habitat of subalpine fir. In forests of Douglas-fir, western white pine (Pinus monticola), lodgepole pine (P.contorta), and even Engelmann spruce, Pacific silver fir is the species that regenerates at greatest rates so as to ultimately replace (to large extent) the other conifers in much of the Northern Cascades, especially on the East Slope. That condition was featured prominently in this photograph where saplings of silver fir formed a "doghair" mid-layer in the vegetation.
Some of the larger saplings or small pole-sized young trees in the "doghair" layer were Engelmann spruce (readily spotted by the reddish hue of their boles), but a high proportion of these had died (as had some of the Pacific silver fir). Death of these smaller trees could have resulted from extreme competition for light (and/or other limiting factors).
At the edge of a small opening in this forest (foreground) various species of shrubs, forbs, grasses, and sedges got enough light to make a last stand before Pacific silver fir filled in most of the remaining forest gaps.
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer), early estival aspect. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Foorest biotic community (!22.5). North Cascades- Okanogan Pine/Fir Hills Ecoregion, 77e (Environmental Protection Agency, undated).
95. Forest forb by the big conifer- Another "photo-quadrant" showing species composition and structure of the cool temperate North Cascades conifer forest considered in this section. Broadleaf or big-leaf lupine was growing at the base of an old-growth Douglas-fir in a local opening while saplings of Pacific silver or Cascades fir dominated regeneration of conifers. Reproduction of Engelmann spruce and Douglas-fir was nil in this forest community.
This photograph also provided an example of the orangish-brown bark of Douglas-fir exposed to full sun for prolongued periods in contrast to the gray or darker-brown color of Douglas-fir bark grown primarily in shade or indirect light (as for example the two trunks presented above).
96. Broadleaf or bigleaf lupine (Lupinus latifolius)- Details of the dominant forb in understorey of the Douglas-fir-Engelmann spruce-Pacific silver fir cool temperate forest described above. This species was described by Hermann (1966, ps. 107-108) as a tall perennial (up to yard tall) and often "...considered one of the best cattle and sheep forage plants" though it can cause sheep poisoning. On many ranges broadleaf lupine apparently is not grazed. The lupones are nodulated, nitrogen-fixing legumes.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer). Peak bloom phenological stage.
97. Another species of shrub at forest edge- Sitka mountain-ash (Sorbus sitchensis var. sitchensis) was the tallest shrub in the Douglas-fir-Engelmann spruce-pacific silver fir forest described above. This member of the rose family (Rosaceae) joined tobacco-brush, bearberry manzanita, and pinemat manzanita as major shrubs that made up two distinct though discontinuous layers in this forest range vegetation. All of these shrubs (and most herbaceous species) were limited to openings or local spots that were devoid of Pacific silver fir which grew so thick as to exclude about everything except the Very Tolerant silver fir.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer).
98. Sitka mountain-ash- Two more views of Sitka mountain-ash showing general features of leaves and inflorescences. There are several mountain-ashes (Sorbus spp.) in the forest of western North America where these shrubs are some of the more distinctive plants below the forest canopy. Sorbus is in the Pomoideae (= Maloideae= Pyroideae) subfamily, the taxon of the classic pome fruits including apples, pears, and two other major rose shrubs of serviceberry (Amelanchier spp.) and hawthornes (Crataegus spp.) which are also important on Northern Cascades ranges.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (early summer); full-bloom stage of phenology.
At similar elevations in the Northern Cascades there are frequently pronounced differences in natural vegetation due to variation in topographic features such as steepness and aspect of slope, exposure, soils, and past and/or current disturbances--or the lack thereof--(eg. fire, flood, blow-down, logging, drought, heavy grazing). In a small, moist valley (a draw) and at slightly lower elevation than that of the preceding Douglas-fir-Engelmann spruce-Pacific silver fir-dominated forest there was a more mesic forest that had developed with a different spceies composition. This forest was dominated by Engelmann spruce that appeared to have persisted as an early seral species and that was being replaced in on-going plant succession by some subalpine fir (Abies lasiocarpa) and, even more, Pacific silver fir, and, most conspicuously of all, western red-cedar (Thuja plicata). There was limited regeneration of Engelmann spruce while most reproduction--thus, ultimate replacement--was western red-cedar, Pacific silver fir, and (though less) subalpine fir.
There was multi-layer, shrub-dominated understorey in this forest that was still in the upper reaches of the Canadian Life Zone (Merriam,1889). This forest community was still on the East Slope of North Cascades (= east of the true Cascade Crest; Matthews, 1999, ps.7-8). It was in the Abies amabilis zone of Franklin and Dryness (1973, ps.93), which means that Pacific silver or Cascade fir is the zonal (=climatic climax) dominant conifer over a a large elevational area (the equivalent of a region in non-monuntainous terrain).
99. Northern Cascades Subalpine Fir-Engelmann Spruce- An old-growth stand of Picea engelmannii that was downslope from the forest just described that consisted of old-growth Douglas-fir (as a persistent colonizing species)-Engelmann spruce with Pacific silver fir and subalpine fir succeeding them. This old-growth Engelmann spruce-dominated forest was contiguous with (adjacent to) the Douglas-fir-Engelmann spruce-Pacific silver fir cool temperate forest described immediately above. The old-growth Engelmann spruce stand now described had developed in a moist depression (draw was probably the most descriptive term), but like like Douglas-fir Engelmann spruce was a seral, early colonizing species that persisted into the climax which ultimately will be dominated by subalpine fir, the major species that was succeeding Engelmann spruce. There was also some Pacific silver fir regeneration, but the more conspicuous successional conifer in moist habitats in this stand was western red-cedar.
The most important difference between this more mesic lowland forest and the more upland the Douglas-fir-Engelmann spruce-Pacific silver fir cool, temperate forest treated earlier was in species of shrubs and structure of shrub layers not in conifer species or the canopy layer. In this old-growth Engelmann spruce-dominated lowland forest there were two shrub layers overall: 1) tall layer consisting of (in wetter habitats) red alder (Alnus rubra), Sitka mountain-ash , red-osier dogwood (Cornus stolonifera) and snowberry (Symphoricarpos alba) and 2) a lower layer of bush honeysuckle, known also as bearberry huckleberry and black twin-berry (Lonicera involucrata), thinleaf huckleberry (Vaccinium membranaceum), grouseberry or whortleberry (V. scoparium), and vine maple (Acer circinatum), this latter shrub ascended to various heights such that it could be interpreted as being in either or both layers. The only herbaceous species having much cover was broadleaf or mountain arnica (Arnica latifolia). At edge of this stand there were plants of the introduced and now- naturalized orchardgrass (Dactylis glomerata).
The first of these two photographs presented physiogonomy of the old-growth Engelmann spruce stand with almost all conifer regeneration by Pacific silver fir, subalpine fir, and western red-cedar and, hence, replacement of Engelmann spruce by these species. Tolerances of these conifer species was undoubtedly a major phenomenon in this change in tree species. Subalpine fire and western red-cedar are Very Tolerant species whereas Engelmann spruce and Pacific silver fir are Tolerant (Wenger, 1984, p. 6). Sitka mountain ash and thinleaf huckleberry made up a tall shrub layer while black twin-berry, whortleberry, and snowberry comprised most of the lower shrub layer. Eyre (1980, p. 105) reported that western red-cedar often dominates stocking on moist sites like flats and valley floors.
The second photogrph featured the tall shrub layer of the understorey along edge of the moist draw through which snow melt water had recently flowed. Major shrubs in this view were red alder, red-osier dogwood, and some Sitka mountain-ash. The major lower-growing shrub was snowberry.
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June, early estival aspect.FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir),but maybe going to or maybe going to SAF 228 (Western Redcedar). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
100. Green layers- "Photo-plots" along a moist draw or drainage running through an old-growth Engelmann spruce-dominated lowland forest in which Pacific silver fir, subalpine fir and western red-cedar were progressing along the sere to succeed Engelmann spruce and to become climatic climax dominants. This forest range was in the Abies amabilis zone of Franklin and Dryness (1973, ps.93). Featured in these two slides was a tall shrub layer made up of red alder, red-osier dogwood, and thinleaf huckleberry with some Sitka mountain-ash and vine maple (which "split the difference" between the two shrub layers).
Western red-cedar was the conifer with the most conspicuous regeneration. There were less regeneration of Pacific silver fir and subalpine fir in the forest range vegetation presented in these two photographs. All large, mature trees were Engelmann spruce.This change in reproduction and the on-going, eventual replacement of Engelmann spruce as subclimax dominant by western red-cedar, Pacific silver fir,and subalpine fir as climax dominants was consistent with the description in Eyre (1980, p. 86) and Johnson and O'Neil (2001, p. 31). Plurality of regeneration stocking on this moist site was typical of western red-cedar (Eyre, 1980, p. 105). Tolerance probably explained much of the shifts in conifer species. Wenger (1984, p. 6) rated western red cedar and subalpine fire as Very Tolerant while Engelmann spruce and Pacific silver fir were rated as Tolerant.
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June, early estival aspect.FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir) or, perhaps,progressing to or maybe going to SAF 228 (Western Redcedar). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
101.Inside a changing old-growth forest- Interior of the old-growth Engelmann spruce-dominated, cool, temperate Northern Cascade forest that was introduced two sets of photographs above. This forest site was more moist (mesic) than forest vegetation on contiguous (=adjoining) sites like the Douglas-fir-Engelmann spruce-Pacific silver fir forest presented immediately before this lowland Engelmann spruce forest. This forest range site was in a small "mini-valley" (more like a draw) that included ephemeral drainage (of snow melt water). The difference in the Douglas-fir-Engelmann spruce-Pacific silver fir forest vegetation described above and this Engelmann spruce-dominated forest that was going to western red-cedar, Pacific silver fir, and subalpine fir was more in the shrub layers (species composition and structure thereof) than in conifer species, execpt for apparent change in dominance of tree species.
Both of these forest range communities were cool, temperate North Cascades forests There was almost no regeneration of Engelmann spruce in vegetation of these forests. In the lowland, subclimax--yet apparently old-growth Engelmann spruce forest--described now western red cedar, Pacific silver fir, and subalpine fir (in that order) were gradually replacing Engelmann spruce. This, as cited earlier, followed successional patterns and adaptations of species described in Eyre (1980; p. 86, 105) and Johnson and O'Neil (2001, p. 31). A large, old-growth specimen of western white pine (Pinus monticola) along with saplings of western red-cedar were consistent with the SAF description of the Engelmann Spruce-Subalpine Fir forest cover type (Eyre, 1980, p. 86). in which western white pine and western red-cedar were listed as major associates in the Engelmann spruce-subalpine fir type in lower and middle elevations in the Cascades.
These two photographs featured the lower shrub layer in contrast to the upper shrub layer presented in the two immediately preeding slides. Major lower-growing shrubs included whortleberry or grouseberry, bush honeysuckle (known also as bearberry huckleberry and black twin-berry), and, less frequently, common snowberry and vine maple. Herbaceous species were limited to non-existent, but the most abundant one was the forb, broadleaf or mountain arnica.
Most of the young conifers (both saplings and large seedlings) representing regeneration/replacement in these two slides was of Pacific silver fir in contrast to earlier-shown slides with western red-cedar as the dominant regenerating species.
East Slope of Northern Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June, early estival aspect.FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir) or, perhaps, progressing to or maybe going to SAF 228 (Western Redcedar). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
102. Forest vegetation stratification- Species composition and structure of the moist, lowland, old-growth Engelmann spruce-dominated forest described in preceding photographs at farther distance from an ephemeral drainage channel. The upper (tall) shrub layer presented previously was absent from this less mesic habitat. With less available soil moisture low-growing shrubs (a low shrub layer) comprised all of the understorey except for cover of regenerated conifers. Regeneration represented in these two photographs consisted of western red-cedar and Pacific silver fir, the two conifer species with most reporduction. ubpresented was the only. Dominant shrubs were grouseberry or whortleberry and bearberry huckleberry (known also as bush honeysuckle or black twin-berry).
This pair of photographs provided a more complete perspective of range vegetation inside this old-growth, lowland forest. A pair of Engelmann spruce trunks served as the focal point of these two photographs (center foreground of first photograph; far left foreground of second photograph). This focal point was retained in the next two-slide set that described the lower shrub layer of this forest range vegetation.
A characteristic of many old-growth forests such as that featured here was the jumble of down timber, all the large logs of which in this forest were Engelmann spruce. Covering country horseback in fallen timber like that shown here is slow to almost impossibe. This forest vegetation is better range for wildlife, even big game species, than for livestock. This was another application of one fo the Four Cardinal Principles of Range Management: Proper Kind and Class of Range Animal.
East Slope of Northern Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June, early estival aspect.FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir) or, perhaps, progressing to or maybe going to SAF 228 (Western Redcedar). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
103. Lowest layer- Two "photo-plots" of the lowest (low-growing shrub) layer of range vegetation in an old-growth Englemann spruce-dominated lowland forest on East Slope, Northern Cascades. This two-slide set provided a summary view (first slide) and a more detailed view (second slide) of the ground- and just-above-ground levels of range vegetation complete with needles and cones of Engelmann spruce along with whortleberry (= grouseberry), the lower of two major shrub species, and bush honeysuckle (= black twin-berry), the taller of the two major shrubs. In the background of both photographs (by spruce log) there were some plants of Sitka mountain-ash. This lower of two shrub layers was the lowest layer of vascular plants in this Engelmann spruce-dominated, lowland cool, temperte forest.
Tree trunks were Engelmann spruce. The two larger, prominent trunks were the focal point in the two photographs that immediately preceded this two-slide set. Together these two two-slide sets provided the reader with a progressive walk-up-to-it view from perspective of a browsing range animal.
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June, early estival aspect.FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir) or, perhaps, progressing to or maybe going to SAF 228 (Western Redcedar). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
104. Another part of the lowest layer- Standing "right next to" a part (say, a zone) of the lower lower shrub layer that was dominated by whortleberry or grouseberry was a zone that was dominated by bush honeysuckle or black twin-berry (foreground). This ericaceous shrub is widespread and common to several forest range communities in the North Cascades and adjoining areas or ecoregions. Other shrub species in background included Sitka mountain-ash, snowberry, vine maple, and thinleafed huckleberry.
Bush honeysuckle was featured in a short series of photographs below. Some of these photographs were taken in the understorey of a black cottonwood (Populus trichocarpa) bottomland forest in early morining of the day after this slide and the other photographs of this old-growth Engelmann spruce-dominated lowland forest were taken.
East Slope of Northern Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June, early estival aspect.FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir) or, perhaps, progressing to or maybe going to SAF 228 (Western Redcedar). Closest classification unit in Brown et al. (1998, p. 37) was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
105. The woods suckle honeysuckle- Bush honeysuckle, twin bearberry, or black twinberry (Lonicera involucrata) is a widespread, often dominant low shrub in forests of the Northern Cascades, including foothills, and forest that develop along streams throughout the Okanogan Upland Region.This stand was growing in the understorey of a Douglas-fir-Engelmann spruce forest at mid-elevation in the Northern Cascades.
Okanogan National Forest, Okanogan County, Washington. June (early summer).
106. Bush honeysuckle blooming - Shoot of bush honeysuckle at peak bloom (first photograph) and details of bush honeysuckle infloresecence (second photograph). These examples were in the colony of bush honeysuckle or black twinberry in understorey of a mid-elevation Douglas-fir-Engelmann spruce forest in the Northern Cascades.
Okanogan National Forest, Okanogan County, Washington. June (early summer).
107. Bush honeysuckle bearing fruit- Bush honeysuckle or black twinberry in understorey of a black cottonwood floodplain forest along the Methow River. Specimens shown here were photographed the day following photographs of this species in the understorey of a Douglas-fir-Engelmann spruce forest at a mid-elevation in the Northern Cascades range. Onset of spring at lower elevations enabled this species to bloom and bear fruit earlier than members of the same species growing at greater elevation in a conifer forest. This was a good application of Hopkin's Bioclimatic Law.
Floodplain of Methow River, Okanogan County, Washington. June (early summer).
At elevations of the general upper montane forest to lower subalpine zones Pacific silver fir becomes the zonal dominant and defining forest species. This is the Abies amabilis zone of Franklin and Dryness (1973, p.93-100). This climax vegetational zone coincides closely with lower to mid levels of the Hudsonian Life Zone (Merriam, 1889). The series of slides presented in the following section dealt with this zone in the interior of the Northern Cascades Mountains. These photographs were taken either from or in the near vicinity of Washington Pass (elevation of 5477 feet) on the third full day of summer (snow banks and all).
108. Vegetation classifier's worry- This Northern Cascades mixed conifer cool temperate or subalpine forest was a transition forest and one that did not fint neatly into any cover type pigeon hole. Dominance of this forest varied locally but often tended to be subalpine fir (Abies lasiocarpa) or, in other local areas, Pacific silver= Cascades fir (A. amabilis). Most regeneration appeared to be A. amabilis, but there was considerable reproduction of A. lasiocarpa. This forest was clearly in the Abies amabilis Zone of Franklin and Dryness, 1973, pgs.93-100), but much of this forest was dominated by subalpine fir that was readily distinguished by the church steeple spire. There was also some lodgepole pine (Pinus contorta) and Engelmann spruce.
The Picket Range of the Northern Cascades creates a rain shadow so that climate is a transition between coastal maritime and interior continental climates, hence a transition forest. The main shrub in much of this understorey, especially in local habitats of snowmelt pools, was the scrub willow known variously as diamond-leaf, flat-leaved, or planeleaf willow (Salix planifolia= S. phylicifolia ssp or var. planifolia). Other important shrub species were thin-leafed blueberry or huckleberry known also as, big huckleberry or tall bilberry (Vaccinium membranaceum) and Alaska huckleberry or Alaska bilberry (V. alaskanense).
Herbaceous species were scarce to absent in this understorey other than for a rare plant of Carex "breeding".
The prominent geologic landmark was Liberty Bell Mountain, elevation 7720 feet.
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
109. The "big picture" and part of the "bell"- Physiogonomy and structure of a Pacific silver fir-subalpine fir forest on the East Slope of the Northern Cascades. Stucture of this simple composition highlands forest was determined primarily by the two principal coniferous species and an even more simple understorey comprise primarily by diamond-leaf or planeleaf willow with sporadic cover of thinleaf huckleberry and/or Alaska bilberry. Snow remaining on the second full day of summer went a long way toward explaining sparsity of an understorey (other than in isolated areas). Pacific silver or Cascades fir was frequently to agenerally the sole dominant of this forest community with subalpine fin the associate. Rarely were the two fir species co-dominants. There were occasional individuals of Engelmann spruce and lodgepole pine, but these were scarse.
Lower layer(s) of woody plants were mostly reproduction of the two fir species (with exception of local assemblages of Vaccinium and Salix species as noted in the preceding paragraph). There were widely scatterred plants of various sedges (mostly Carex spp.).
Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). In the Hudsonian Life Zone (Merriam, 1889). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
110. The next generation in a local clearing- A small opening in the canopy of a climax old-growth (or approaching old-growth stage) Pacific silver fir-subalpine fir forest that once existed as a local "micro-meadow" sedge (mostly Carex spp.) and tall or manysided cottongrass (Eriophorum polystachion= E. angustifolium) being invaded as a typical "doghair" stand of larger seedlings of Pacific silver fir. Pacific silver fir was the overall dominant tree with subalpine the associate species at elevations somewhat below its "prime habitat". At edge of this small natural clearing there mature and old-aged trees of both fir species though Pacific silver fir predominated.
Note the disheveled or tossed-about appearnce fo Pacific silver fir seedlings caused by heavy accumulations of snow (eg. some still present immediately to left of the partially bent-over seedlings).
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). In the Hudsonian Life Zone (Merriam, 1889). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
111. Stand structure and a staid composition- "Broadside" view of a climatic climax highlands forest dominated overall by Pacific silver fir with subalpine fir. There were some individuals of lodgepole pine and Engelmann spruce. There was sporadic cover of an understorey made up mostly of planeleaf or diamondleaf willow with small, localized patches of huckleberry species. This was an example of the often-found phenomenon where species diversity is low in climax vegetation.
The tolerance ratings of Tolerant and Very Tolerant for Pacific silver fir and subalpine, respectively, (Wenger, 1984, p. 6) were obvious in this photograph that showed abundant regeneration of both Abies species.
At the great distance from which this photograph was taken (to present a "feel" for physiogonomy and structure of this cool, temperate highlands fir forest) even Abies saplings were inconspicuous. Diamondleaf willow (and probably other Salix spp. as well) were represented only as brown spaces in the understorey. At this season or point in the annual growing cycle of deciduous species willow buds were not yet open so that the presence of willow was marked by the color of fir needles and not the characteristic green shades of willow leaves.
This natural (climatic climax) vegetation has generally been interpreted as subalpine rather than a montane forest and as a transition forest between coastal maritime and interior continental climate due largely to the rain shadow affect created by the Picket Range of the Northern Cascades.
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). In the Hudsonian Life Zone (Merriam, 1889). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
112. More valuable for water than forest or range commodities- A localized, small snowbank still present on the second full day of summer indicated the importance of the Pacific silver fir-subalpine fir forest as watershed, including provision of melt water for wildlife and fisheries, irrigation, municipal uses, aesthetics, recreation, etc. Recreational uses of snow would include hunting, wildlife watching, and cross-country sking. In dense (even "doghair" ) fir stands like those shown in this series of photographs cross-country sking would be problematic if not impossible. Steepness of slopes does not help matters. Value of high-elevation forests (including mountain hemlock and subalpine fir described below) for retention of snowpack for irrigation water and flood control cannot be overemphasized.
This forest vegetation was made up mostly of Pacific silver fir and, at lesser cover, subalpine fir, including the lower layers (those below tree canopy) which consisted primarily of Abies regeneration except for small openings where willow, bluebery (huckleberry), and sedge species were important. Cimax (climatic) status of this forest was obvious from this uneven-aged structure that showed reproduction of these two species in or under "their own shade".
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). At elevations of the Hudsonian Life Zone (Merriam, 1889). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
113. Interior of a Pacific silver fir forest- Structure of a zonal (= climatic climax) pacific silver or Cascades fir-dominated highlands forest various layers of forest range vegetation well-represented. This included the downed timber or fallen log component that is characteristic of most old-growth or near old-growth conifer forests.
Several diffrent age classes of Pacific silver fir were obvious in this "photo-quadrant" including seedling, sapling, and pole size categories. It was a conspicuously uneven-aged forest stand. This age clsss-structure was featured to highlight natural regeneration of this climax conifer and to "photo-document" the tolerance rating of Tolerant for Pacific silver fir (Wenger, 1984, p. 6). Although subalpine fir was rated as Very Tolerant (in this same reference) there was usually much less of this latter than of the former fir species in this cool, temperate, transition forest.
Also present--though not conspicuous--were willows, especially diamondleaf or flatleafed willow, thinleafed blueberry (huckleberry) and Alaska bilberry (huckleberry), and scattered sedges of unknown species. These angiosperm species were still dormant (or under snow).
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). At geleral elevations of the Hudsonian Life Zone (Merriam, 1889). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
114. Local co-dominants and an understudy- Bottom portions of trunks of subalpine fir (Abies lasiocarpa) on the left; Pacific silver or Cascades fir (A. amabilis) on the right. These were the climatic climax dominants of the highlands transition fir forest treated in this part of the publication. A lower woody layer made up of planeleaf or diamondleafed willow was present in the right corner of this "photo-plot". Buds on the willows were shown in swollen later stages just prior to opening for their summer growing season. (Photograph was taken on second full day of summer.) Locally there were species of huckleberry or blueberry, especially thinleafed huckleberry and Alaska bilberry.
Willow buds are an important feed source to white-tailed ptarmigan (Lagopus leucurcus) which are year-round residents of the North Cascades.
The high degree of tolerance in both of these species was apparaent from the numerous fully leafed, lower (almost to the ground) boughs.
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). At geleral elevations of the Hudsonian Life Zone (Merriam, 1889). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
115. Another Pacific silver fir-subalpine stand on a more exposed slope- At a location different from the one treated immediately above a similar Pacific silver fir-aubalpine fir-dominated highlands forest had developed with about the same species composition and vegetational structure. This latter forest was on a more northerly slope and a more mesic (almost wet) site due to presence of more (or later arrival of) melt water.
Lower layers of forest vegetation were similar with dominant or more common shrub species being planeleaf willow and various huckleberry species (Vaccinium membranaceum, V. alaskanense). Various age classes of both Abies species were present from the low shrub layer (seedlings and small saplings) to mid-height (large poles) up to the canopy layer. Herbacaeous species such as sedges were nominal except in small to tiny forest openings.
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). At general elevational zone of the Hudsonian Life Zone (Merriam, 1889). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
116. Inside of another Pacific silver fir-subalpine fir stand- Two photographs that portrayed plant species composition and structure (layers) of vegetation of this forest range. The first slide was a closer-in view of the "photo-plot" introduced in the preceding photograph (and that was marked--however fleetingly--with a small patch of snow on this second day of summer). The second slide was another sample of this forest's range vegetation. The low shrub layer of willow (mostly planeleaf willow) and scattered plants of thinleaf and Alaska huckleberry was present and obvious in both of these "photo-plots". Herbaceous plants were largely absent throughout the area of a Pacific silver fir-subalpine fir forest presented here.
The unevern-aged structure of this stand, like the one discussed just prior to it, showed the Tolerant and Very Tolerant rating of Pacific silver fir and subalpine fir, respectively (Wenger, 1984, p. 6). This in turn attested to the climax nature of this forest range vegetation. Most angiosperms except for the Vaccinium species were still in dormancy (or under snow cover). Even the huckleberry species were "a while away" from blooming.
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). At general elevational zone of the Hudsonian Life Zone (Merriam, 1889). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-3 (Silver Fir-Douglas-fir Forest). No appropriate SAF designation because there was not an SAF forest cover type of Pacific silver fir. Brown et al. (1998, p. 37) unit of vegetation classification was Pacific Silver Fir Series (122.54) of Cascade-Sierrean Montane Conifer Forest biotic community (!22.5). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
117. Another (and sometimes forgotten) part of the highlands fir forest- In his college course, Rangeland and Forest Plants, this author routinely indicates to his students that what is or constitutes a "range plant" depends on personal interpretation and what sources were consulted. In the old or classic Botany, any living thing that was not an animal was a plant (or vice versa). Bacteria, algae, fungi, lichens, and even viruses as well as bryophytes and vascular plants were all "plants". With advent of additional biotic kingdoms beyond the traditional two kingdom system such as that of Whittaker (1969) the term "plant" (and, to some extent, the study of "plants") was thrown into a tailspin.
Now what is a "range plant"? In their text, Biology of Plants, Raven et al. (1992) did not even include the term "plant" in their glossary! Furthermore, Raven et al. (1992, p. 11) stated that they did not regard algae, fungi, bactreria, or viruses as palnts and would not refer to them as plants, but they included these non-plants in a basic textbook on plants "because of tradition" and, due to convention, these organisms were "part of the botanical portion of the curriculum". They ain't plants, but we put them in our Botany textbook, Biology of Plants, anyway (and we're going to keep Plants in the title of our text that includes organisms that we would not call plants). OK ...
The author of this publication was not so "slick". He simply regarded these non-plant plants as parts (components) of the range vegetation which he sought to present and describe to fellow rangemen and others interested in range cover types. In that spirit two photographs of a slow-melting, remnant snowbank were presented as coverage of the natural range vegetation of a Pacific silver fir-subalpine fir highlands forest.
Red snow. Watermelon snow. A small snowbank photographed on the second full day of summer in the near vicinity of Washington Pass (elevation, 5577 feet) in the North Cascades supporting colonies of snow algae. The red or reddish brown coloration of this "dirty snow" was due to any of several species of algae. In discussing watermelon snow Mathews (1999, p. 482) remarked that over a hundred species of snow algae had been identified. The most common of these is Chlamydomonas nivalis, a single-celled motile green alga and specifically a flagellate (with reference to the flagellum which is the thin organelle that enables one-celled organisms to move).
This is a good example of a protist (Allaby, 1998) which in this species combines animal-like locomotion and an eyespot (Wilson et al., 1971, p. 458) with plant-like cellulosic cell walls and chlorophyll so as, according to Mathews (1999, p. 482), preclude this organism from being labeled either plant or animal. In C. nivalis there are carotenoid pigments that produce the red color color in this green algae which helps protect these single-celled organisms from photodestruction that would otherwise occur due to the intense light striking slowly melting snowbanks (Ray et al., 1983, p. 441). Standard textbooks in basic Botany routinely devoted some space to Chlamydomonas species. Interested students were directed especially to Arnett and Braungart (1970, ps. 6, 244, 354, 355) and Wilson et al. (1971, p. 458) as well as the just-cited Ray et al. (1983). Transeau et al. (1953) used a photograph (plate 1) of red snow as half of the frontispiece of their their classic text, Textbook of Botany, which showed the appreciation these authors had for simple, nonvascular plants.
Red snow is notarious for presumedly causeing diarrea in thirsty mountaineers so descretion would suggest consuming such colored snow as tea or some other beverage brewed from the snow as boiled water (ie. at least boil the water briefly).
In final summary, algae like lichens, fungi, and so on are part of the range vegetation: in the high forest, out of the forest, in a snowbank, whatever. What, if any, benefit or detriment snow algae have on range animals (other than rangemen) was not known to this author.
East Slope of North Cascades, Methow Valley Ranger District, Okanogan National Forest, Okanogan County, Washington. June (the second full day of summer), late vernal aspect. Washington Pass (elevation of 5477 feet). At general elevational zone of the Hudsonian Life Zone (Merriam, 1889). Part of the vascular plant-defined range vegetation whose ecosystem, Kuchler unit, forest range cover type (lack thereof), biotic community, and ecoregion were given immediately.
North Cascades subalpine meadows and tree groups- Westward and downslope from Washington Pass (elevation of 5477 feet) is Rainy Pass (4855 feet). This was the highest part of the Hudsonian Life Zone (Merriam, 1889) in this part of the Northern Cascades being just below timberline or start of the Alpine Life Zone. The following set of photographs was of subalpine parklands including subalpine mountain meadows dominated by black alpine sedge (Carex nigricans), cotton-grass, primarily tall or manysided cotton-grass (Eriophorum polystanchion) and sphagnum mosses (Sphagnum squarrosum, S. magellanicum)---the Eriophorum- Sphagnum Association---being invaded by conifers, often exclusively by subalpine fir. This phenomenon was described in Franklin and Dryness (1973, ps. 108-109, 272-284). There were some other carices like water sedge (Carex aquatilis) that were locally dominant. The most conspicuous and, usually, most common shrub on subalpine meadows---aside from avalanche chutes---was flat-leaved = planeleafed willow (Salix phylicifolia var. planifolia= S. planifolia).
Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). North Cascades- North Cascades Subalpine/Alpine Ecoregion, 77c (Environmental Protection Agency, undated).
118.The difference clouds (overcast) made- Two paired photographs taken just a few minutes apart of a small valley and mountain crest in North Cascades in mid-afternoon of early summer. In the first first photograph there was a generally overcast sky with light (less dense) cloud cover at heights of mountain peaks. In the second photograph there was still some overcast but less cloud cover and low clouds previously at heights of mountain tops (ie. cloud floor down to the Nival Zone) had lifted. This pair of slides was presented so as to compare (or contrast) natural lightening to illustrate the difference in color photography that is frequently produced by cloud cover. In this comparison (contrast) there was not doubt but that cloud cover at this elevation resulted in two different conditions of light and different appearance of vegetation and geologic features. For most purposes less cloud cover resulted in a superior (more color contrast and detail at distance) photograph although some detail at this distance of trees and tree groups were more clear under (with) more (heavier or denser) cloud cover. Such is not always the outcome, but it usually is, especially with heavy and/or low cloud cover. At higher elevations where there is much greater ultraviolet radiation, some cloud cover-- light (vs. heavy or dense) with high clouds--typically results in a naturally "UV light-filtered" effect resulting in more sharply contrasting colors. This was shown here where details of dark trees and groups of trees were superior under low cloud cover that was not heavy or dense cloudiness (ie. some but not too much overcast). Conversely whiteness of snow was brighter, clearer, and more distinct under more not less sunlight. Atmospheric conditions that are ideal for some photographic purposes are sometimes substandard for other pictorial purposes.
Now for the range vegetation (the purpose of the photographs). Subalpine mountain meadows being invaded primarily by subalpine fir with some Engelmann spruce and lodgepole pine and, infrequently to rarely, mountain hemlock.Mountain hemolock-dominated forests were "just down the hill" from this right-below-timberline mosaic of North Cascades range vegetation. In addition to the Hudsonian Life Zone featured in these photographs the patchwork of forest, meadow, alpine, snowfield, and "rock and ice" vegetation throughout this panarama included the Alpine and Nival Life Zones (Merriam, 1889) higher up in the mountains (and farther back in the photographs).
The dynamics of forest and tree groups and subalpine mountain meadow was emphasized in photographs presented immediately below. While a case could be made--especially froma rangeman's perspective--that most of this potential natural range vegetation was meadow not forest, the present or existing vegetation was more that of tree-dominated range communities. For that reason this range vegetation was included with forests of the Northern Cascades and not in the mountin meadows section (Grasslands- Meadows). Such ecotonal vegetation (between subalpine forests below and alpine meadows above) constitutes timberline range communities. Franklin and Dryness (1973, ps. 108-109, 272-284) remained the classic treatment of this transition zone range vegetation.
Landscape-scale perspectives of this North Cascades landscape was noted immediately below where the application and usefulness of Landscape (vs. Ecosystem) Ecology was remarked.
East Slope of North Cascades, Rainy Pass (4855 feet). Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (the third full day of summer), late vernal aspect. Natural range vegetation of foreground (assuming plant community was primarily forest and excluding Alpine and Nival Zone communities from consideration) was most closely classified by the following units. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock). SAF 206 (Engelmann Spruce-Subalpine Fir) mostly; some SAF 205 (Mountain Hemlock). Surprisingly there was no appropriate unit given by Brown et al. (1998, p. 36-37) because this would be a Subalpine Fir Series under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4). North Cascades- North Cascades Subalpine/Alpine Ecoregion, 77c (Environmental Protection Agency, undated).
119. Losing the mountain meadows- Subalpine mountain meadows (what there was left of them) under on-going massive invasion by subalpine fir with much less cover and density of Engelmann spruce, lodgepole pine, and, least, mountain hemlock. This successional phenomenon of conifer invasion was dealt with by Franklin and Dryness (1973, ps. 108-109, 272-284) to which readers were referred. The complex and interacting set of ecological variables responsible for this woody invasion into subalpine meadows (which are part of the potential natural vegetation of this mosaic of mountain plant communities) is incompletely understood and beyond scope of this publication. Photographic examples (and further documentation) of the dynamics of contiguous range plant communities was deemed sufficient for this pictorial purpose.
Important range plant species of meadows included cotton-grass, primarily tall or manysided cotton-grass, sedges like water sedge, and sphagnum mosses. Meadow vegetation preented here and in succeeding photographs was mostly a combination of the black alpine sedge (Carex nigricans) community and the Eriophorum- Sphagnum Association (Franklin and Dryness, ps. 108, 261). There were also clumps of different willows (Salix spp.) of which flat-leaved = planeleafed willow was most common and important. It was explained above when describing Pacific silver fir-subalpine fir forests that flat-leaved willow was a major shrub species in the montane and subalpine forests of the North Cascades.
These two photographs provided a landscape-scale view of the greater North Cascades landscape illustrating the utility of certain concepts of Landscape Ecology. It was obvious that Ecosystem Ecology--in fact, the very concept of ecosystem--had limitations when the lens of a 35 mm camera at human eye-level captured so many diverse ecosystems (subalpine conifer forest, willow stands, mountain meadows, avalanche paths, snowbanks, talus slopes, and rock mountain domes) in three Life Zones (Hudsonian, Alpine, Nival). Fortunately range and forest ecologists and practitioners can benefit from both subdisciplines or special fields of Ecology.
East Slope of North Cascades, Rainy Pass (4855 feet). Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (the third full day of summer), late vernal aspect. Natural range vegetation of foreground (assuming plant community was primarily forest and excluding Alpine and Nival Zone communities from consideration) was most closely classified by the following units. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock). SAF 206 (Engelmann Spruce-Subalpine Fir) mostly; some SAF 205 (Mountain Hemlock). Surprisingly there was no appropriate unit given by Brown et al. (1998, p. 36-37) because this would be a Subalpine Fir Series under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4). North Cascades- North Cascades Subalpine/Alpine Ecoregion, 77c (Environmental Protection Agency, undated).
120. Crowding in, closing off, and losing out- Subalpine mountain meadows and subalpine conifer forests naturally exist in a state of dynamism and successional tension with relation with each other. Fires, drought, snowpack (depth of snow accumulations), grazing/browsing, windthrow (= blowdown), insect and disease outbreaks, etc. create constant ebb and flow, advance and retreat of first one and then the other of these two natural range plant communities at subalpine elevations in western mountains that have prolonged cool or cold temperature regimes. A patchwork of these two major range types--plus smaller local stands of conifers known as tree groups along with incidental range plant communities like those along streams (riparian communities) and avalanche troughs--results in a complex of ecosystems and local environments interacting at landscap-scale as seen here in the interior of the Northern Cascades.
Over course of the last century or so the successional "momentum" has been in direction of conifer invasion (encroachment) into the subalpine meadows and expansion of tree groups which further reduced size and scope of mountain meadows. This is, of course, not possible at higher elevations of alpine range which, by self-evident definition, is above timberline. At elevations below timberline (subalpine and montane; Hudsonian, Canadian, Transition Life Zones) woody plants can appear, disappear, and reappear indefinitely with march of time and plant succession along these various seres. Throughout many of the higher elevation (or colder temperature) mountain ranges at subalpine elevations where the alpine meadows and subalpine forests are engaged in a constant tug of war for survival this vegetational dynamic has had a dramatic and overwhelmingly one-sided shift to conifer cover and resultant loss of subalpine meadows.
This dramatic shift in plant species and eventual change in type of range plant community was obvious in these two photographs. Conifer seedlings and small saplings (most of subalpine fir) had invaded even into the innermost parts of wet meadows. A small meandering stream was on the range shown in the first of these two slides. A small, round-bottom valley with some surface melt water was portrayed in the second slide. Herbaceous vegetation of what remained of the subalpine mountain meadows was largely of the black alpine sedge community and the Eriophorum- Sphagnum Association (Franklin and Dryness, 1973, ps. 108, 261), with black alpine sedge and tall or manysided cottongrass the dominant or associate grasslike plant species depending on local habitat. There were probably also some other kinds of sedges, but all Carex species were in half-rotten vegetative states making almost all plant identification impossible for visitors to these ranges.
Conifers were overwhelmingly subalpine fir (a consociation of that species) but there were numerous--usually widely scattered--individuals of Engelmann spruce (especially closer to moist soil conditions), lodgepole pine, mountain hemlock, and, least of all or rarely, Paccific silver fir. Invasion of subalpine mountain meadows begins a successionally cascading effect in which deeper-rooted trees absorb and transpire more water from the soil that in turn results in continued drying out of soil which then makes the drier environment even more "inviting" to more conifes which prefer better-drained soils. Eventually a former natural wet mountain meadow becomes a subalpine conifer forest.
What prevented this shift in range type in the past? Or, alternatively, has this phenomenon repeatedly taken place on a successional time scale in the ecological past with dynamic wetting-and-drying of subalpine soils? Is this change in range vegetation merely the last "conifer part of the cycle" with trees currently having their successional day in the sun? Or is this change from herbaceous vegetation to woody vegetation (both types comprised of native species) an unnatural developmnent initiated and/or maintained by human induced disturbances perhaps including change in fire regime (ie. reduction of even elimination of fire or, alternatively, too many wildfires), overgrazing (either past or present, livestock and/or wildlife), undergrazing, or even local or planetary climate change?
Incidentally, these two photographs were taken under a full-sun sky (scattered cloud cover was present beyond the land photographed here) so that there while there were some shadows cast there was overall more detail and a superior view of range vegetation.
East Slope of North Cascades, Rainy Pass (4855 feet). Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (the third full day of summer), late vernal aspect. Natural range vegetation of foreground (assuming plant community was primarily forest and excluding Alpine and Nival Zone communities from consideration) was most closely classified by the following units. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock). SAF 206 (Engelmann Spruce-Subalpine Fir) mostly; some SAF 205 (Mountain Hemlock). Surprisingly there was no appropriate unit given by Brown et al. (1998, p. 36-37) because this would be a Subalpine Fir Series under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4). North Cascades- North Cascades Subalpine/Alpine Ecoregion, 77c (Environmental Protection Agency, undated).
121. A last look, but what hopefully will not be the last look, at a North Cascades wet mountain meadow- A fine example of subalpine, wet mountain meadow surrounded by subalpine conifer forests and tree groups. These stands were dominated by subalpine fir with some trees of Engelmann spruce, lodgepole pine, and mountain hemlock (even a few Pacific silver fir). Conifers had invaded and were continuing to invade the wet meadow.
A small, widely meandering stream flowed through this subirrigated wet meadow that was comprised of black alpine sedge, tall cottongrass, other sedges (mostly Carex spp), and sphagnum mosses. Meadow vegetation was mostly of the black alpine sedge-dominated community and the Eriophorum- Sphagnum Association (Franklin and Dryness, 1973, 108, 261).
Invasion followed by aggregation of the deeper-rooted conifers could ultimately result in loss of many or, even, all meadows due to drying of wet meadow soils as trees absorb and transmire more water. Continual and progressive drying of soils (ie a form of biotic land drainage) results in an on-going cycle in which the environment (especially the edaphic portion thereof) becomes ever more amenable to establishment of conifers whose "prime habitat" involves well-drained soils.
Over periods ranging from about 50 to 125 years there has developed well-established pattern of drying out of meadows and driving out of wetland range plants with establishment of conifers and eventual replacement of meadow with subalpine conifer forest. Will this successional trend continue? If it does, is this the natural state of native plant communities? Is it just part of a larger successional cycle? Have droughts caused by sunspot activity had a role in this change in range vegetration. Or is this phenomenon the result of anthorpogenic action (ie, man's abuse of the land through improper burning, grazing, change in chemical composition of the atmosphere)?
Is this a last look at this meadow or just the most recent stage in the dynamic development of vegetation?
East Slope of North Cascades, Rainy Pass (4855 feet). Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (the third full day of summer), late vernal aspect. Natural range vegetation of foreground (assuming plant community was primarily forest and excluding Alpine and Nival Zone communities from consideration) was most closely classified by the following units. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir-Hemlock). SAF 206 (Engelmann Spruce-Subalpine Fir) mostly; some SAF 205 (Mountain Hemlock). Surprisingly there was no appropriate unit given by Brown et al. (1998, p. 36-37) because this would be a Subalpine Fir Series under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4). North Cascades- North Cascades Subalpine/Alpine Ecoregion, 77c (Environmental Protection Agency, undated).
North Cascades mountain hemlock forest- At elevations lower than the subalpine parklands, westward, and on harsher forest sites (more exposure, shallower and/or drier soils, etc.) in the North Cascades subalpine fir yielded dominance to western hemlock. This is the Tsuga mertensiana Zone of Franklin and Dryness (1973, ps. 101-109). Two examples of subalpine forests (two separate forest tracts) were shown and described in the following section of this publication. In the forest range vegetation presented in the following photographs mountain hemlock was the dominant species with subalpine fir, Engelmann spruce, or western hemlock as the local associate species. Mountain and western hemlocks often grew cheek-by-jowl and with cones of both of these two Tsuga species intermixed on the ground. Engelmann spruce and lodgepole pine were locally common and subalpine fir and/or Pacific silver fir held its own such that there was substantial local (stand-to-stand) variation. Understorey layers were usually limited--especially herbaceous plants, and in the old-growth examples presented here--but important shrub species were primarily those listed for the higher subalpine parklands described above and for the lower elevation, adjoining subalpine-Engelmann spruce type that was described below.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
Subalpine fire western red cedar, and western hemlock are Very Tolerant Engelmann spruce, Mountain hemlock and Pacific silver fir are Tolerantis Douglas-fir an western white pine are intermediate in tolerance (Wenger, 1984, p. 6).
122. General appearance- Physiogonomy, structure, and species composition of a North Cascades (East Slope) mountain hemlock-dominated subalpine forest. Two photographs showing exterior and physiogonomic aspects of a cool-temperate subalpine forest in which most of stocking was mountain hemlock with Engelmann spruce (often the associate species; persisting as large trees from previous seral communities), western hemlock, subalpine fir, Pacific silver or Cascades fir, and even some lodgepole pine "rounding out" the list of major coniferous species. The understorey was largely undeveloped other than locally where adequate light penetrated this otherwise dense canopy forest. Major shrub species included thinleaf, big, or black huckleberry or, sometimes, blueberry (Vaccinium membranaceum) and Alaska huckleberry or bilberry (V. alaskaense). Along small drainages (more meltwater springs or seeps than streams) willow was abundant. Most of this was diamondleaf willow. Herbaceous plants were downright rare with Carex being about the major genus represented.
Along outer edge of this forest there was regeneration of Engelmann spruce, a local associate speices, but there was less spruce reproduction farther inside the forest. There was also regeneration of subalpine fir and even some of Pacific silver fir, but most reproduction overall was of mountain hemlock indicating (or, at least, suggesting) that this was the climax conifer. In the interior of this subalpine forest there was amble regeneration of western hemlock to maintain this as a local associate species (see below, this series of slides). Regeneration of these coniferous species was in line with tolerance ratings provided by Wenger (1984, p. 6) which put subalpine fir and western hemlock as Very Tolerant and mountain hemlock, Pacific silver fir, and Engelmann spruce at Tolerant. In effect, all of these conifers were capable of regenerating their species in this cool, temperate, subalpine forest.
Likewise there was even some lodgepole pine as a minor species. Note the small sapling representing this species in the left foreground of the first photograph.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
123.Closer look at structure and species composition- Another view of the mountain hemlock-dominated subalpine forest introduced above. All of the larger (mature) trees in this "line-up" were mountain hemlock except for the big tree on far left and the big trunk second from right which were Engelmann spruce. While there was some regeneration of mountain hemlock "in its own shade" a disproportionately high proportion of conifer reproduction in the vegetation of this and the near-vicinity of this photograph was of subalpine fir. There was also some--though correspondingly less--regeneration of Pacific silver fir and still yet less reproduction of Engelmann spruce and western hemlock in this "photo-plot".
This was an old-growth or near old-growth stand. There was some regeneration of all coniferous species although reproduction was apparently more abundant.in subalpine fir and mountain hemlock. The most frequently encountered shrubs were thinleafed or black huckleberry and Alaska huckleberry (bilberry) which formed small local "patches" (colonies).
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
124. Big trees high in the hills- Two "photo-plots" of species composition and layers of vegetation in the mountain hemlock-dominated subalpine forest presented immediately above at scale of physiogonomy and structure. In the first "plot" two mature and quite large mountain hemlock stood as "doormen" indicating the dominant conifer (and forest dominant) of this cool, temperate Northern Cascades forest. The second "plot" featured an immense old-growth specimen of Engelmann spruce, a local associate species of this highlands conifer forest. Regeneration at base and to rear of this still-in-its-prime spruce included Engelmann spruce seedlings and saplings, but also subalpine fir and mountain hemlock.
Even though this was a mountain hemlock-dominated forest it was a conifer forest community made up of a mixture of Engelmann spruce, subalpine fir, western hemlock, Pacific silver fir, and even isolated lodgepole pine. Coniferous species from neighboring forests of various cover types, especially Pacific silver fir and subalpine fir-Engelmann spruce, were well-represented in this specific forest. This was particularily the case at spatial scale of local stands.
The understorey was poorly developed in this closed canopy, subalpine forest. Herbaceous species were almost absent so that there was not an herbaceous layer. A sporadic or discontinuous shrub layer was made up primarily of thinleaf, big, or black huckleberry and Alaska huckleberry or bilberry. Willows, most of which were diamondleaf willow, grew along and even in bottoms of small, shallow meltwater drainages.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
125. Close up inside- Interior of the mountain hemlock-dominated subalpine forest forest shown previously at larger spatial scale (sets of slides immediately above).
The affilitation or together-aggregation of mountain hemlock and subalpine fir was recognized by the Society of American Foresters of this forest cover type by the former designation of Mountain Hemlock-Subalpine Fir (Eyre, 1980, p. 85).
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated). .
126. Showing their trunks- Another perspective of the interior of the mountain hemlock-dominated subalpine forest forest presented hererin. Trunks of various sizes (and presumedly of ages) of mountain hemlock was represntative of a conifer that is a climax tree species on this forest range site and of the forest cover type named after this dominant climax species.
Lack of a well-developed understorey--if there were any lower layers of vegetation at all-- in this photograph was typical of this subalpine conifer forest. The Society of American Foresters description of this forest cover type (Eyre, 1980, p. 86) explained that mountain hemlock forests vary from having "deparuperate understories" to "relatively rich mixtures of three to six species...".
Portions of the ground surface and upper horizons of soil had large rocks, even boulders, of igneous intrusive material, including granite.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
127. Beauty in structure- Exterior view of a mountain hemlock-subalpine fir-western hemlock forest in the interior of North Cascades showing physiogonomy and layers of woody vegetation. This photograph was the first in a second series of slides that presented a second subalpine conifer forest dominated by mountain hemlock. Western hemlock, locally the associate species, was more common and accounted for a greater proportion of canopy cover than in the forest tract described immediately above. In this forest subalpine fir, an infrequent local associate species, was less common and had lower regeneration than in the forest described immediately above. Also, Pacific silver fir and lodgepole pine had much less representation, both as to canopy cover and regeneration.
This was a transition zone forest situated below higher-elevation (at highest subalpine zone; immediately below alpine) subalpine fir-dominated forests and above Engelmann spruce-subalpine fir forests that developed in areas of drainage along mountain slopes.
Young conifers (mostly mountain hemlock, western hemlock, subalpine fir, and Engelmann spruce in that order) of various age/size classes formed a well-developed woody layer below that of the forest canopy. This was ecotonal forest vegetation of these various conifer species. Its composition was typical of ecotones in having greater species diversity.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
128.Start of another "look-see"- Interior of the ecotonal mountain hemlock-subalpine fir-western hemlock subalpine forest introduced in the preceding slide. There were mature, pole, sapling, and seedling ages/sizes of all of these species as well as of Engelmann spruce (though this latter species was much less abundant or represented in this forest tract).
Conifer regeneration formed a lower woody layer at considerable distance below the canopy layer. Trees of the various ages/sizes often grew in close proximity to each other (indicative of Tolerant and Very Tolerant species), but this was of such irregularity that the woody understorey did not form multiple layers in the usual sense. Most of the larger trees were mountain hemlock and western hemlock.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
129. From hanging tall to the ground- Structure and species composition of a mountain hemlock-subalpine fir-western hemlock subalpine ecotonal forest. Two views of vertical stratification of the transition, subalpine, conifer forest descriped in the two preceding photographs and captions.
.East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
130. Into the woods for the "look-see" started earlier- Two photographs of the interior of views of a mountain hemlock-subalpine fir-western hemlock subalpine ecotonal forest (the second forest tract of subalpine conifer forest dominated by mountain hemlock described under Northern Cascades).
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated). -
131.A "looksee" at forest structure and composition- Interior of a mountain hemlock-subalpine fir-western hemlock subalpine forest that was an ecotone (transition zone) between a North Cascades Engelmann spruce-subalpine fir forest that had developed in a moist draw at slightly lower elevation and a higher elevation subalpine, wet mountain meadow (parkland) range that was surrounded by subalpine conifer forests dominated by subalpine fir. The subalpine forest vegetation presented in this photograph western hemlock was an associate to co-dominant subalpine fir and mountain hemlock. Both hemlock species often grew side-by-side with cones of these two conifers intermixed on the soil (ground) surface. An example of this was shown at far right of this slide: western hemlock (rightmost tree in photograph) and mountain hemlock (trunk with deeper furrowed bark and immediately to left of the rightmost western hemlock).
Vegetation layers below the tree canopy zone included an upper or higher shrub layer of vine maple (most of which was dying), a lower shrub layer of young huckleberry of undetermined species, the large round leaves of which suggested ovalleaf huckleberry (Vaccinium ovalifolium), and a soil surface layer or zone of moss. The arboreal layer of moss and lichens that was noted in preceding photographs was also obvious in this photographre. Although these epiphytes were in the conifer canopy they constituted what could plausibly be viewed as a separate layer in this forest vegetation.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
132. A last long "look-see"- Local stand of western hemlock (the three big trunks) with abundant regeration of this species, but also of subalpine fir and some (relatively small proportion) of Engelmann spruce. Layers of this forest vegetation were featured in this "photo-plot" including downed logs and species of lichen and true mosses hanging from tree branches. Lichens and bryophytes on conifer branches formed a distinct component of the canopy layer.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-4 (Fir Hemlock Forest). SAF 205 (Mountain Hemlock). Mountain Hemlock Series (121.43) under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) of Brown et al. (1998, p. 36-37). North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
133. Conked on the butt- Lower trunk of an old-growth (and "more-dead-than-alive") specimen of Engelmann spruce in the western hemlock-dominated forest displayed and described immediately above. Several so-called fruiting bodies (reproductive structures) of the polypore fungus, red-belted conk (Fomitopsis pinicola= Fomes pinicola), adorned this old bole on the lowermost butt portion (at union of trunk and upper lateral roots). Some kind moss and various ferns also used bark and microhabitat of this trunk for their dwelling place. Moss was strictly a commensal. Not necessarily so this common, woody conk.
Arora (1986, ps. 578-579) reported that red-belted conk "attacks" (ie. consumes via decomposition) rwood of the host tree (usually it is already dead) resulting in a carbonizing rot (brown crumbly rot) that breaks the wood grain and turns wood fiber brown. This polypore species is a major reducer of dead conifer wood.
This range plant (plant according to the traditional two-kingdom classification which this author retained throughout) provided another example of organisms that are (function as) consumers or reducers in range ecosystems.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer).
Northern Cascades subalpine fir-Engelmann spruce forest- At lower elevations and westward from subalpine meadows-subalpine fir tree groups in the North Cascades and in moist areas such as depressions and stream bottoms subalpine fir-Engelmann spruce forest communities developed often with both western and mountain hemlocks (Tsuga heterophylla and T. mertensiana, respectively) as associate species. There was typically little or no understorey in these forest other than in forest openings and at forest edge. Here the dominant shrubs included thinleaf and Alaska huckleberry or bilberry (Vaccinium membranaceum, V. alaskense), Sitka mountain-ash (Sorbus sitchensis), black elderberry (Sambucus racemosa var. melanocarpa), and common snowberry (Symphoricarpos albus). Herbaceous species were limited and typically non-existent.
This forest provided textbook example of tolerance and plant succession. The Very Tolerant subalpine fir was replacing the Tolerant Engelmann spruce. Somehow both western and mountain hemlocks were present. These Tsuga species were dominants (or associate species) of the higher elevation, more exposed slopes above the moist and more sheltered subalpine fir-Engelmann spruce forest range type. Obviously the moist subalpine fir-Engelmann spruce forest was transitory range.
Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (second full day of summer). North Cascades- Pasayten/Sawtooth Highlands Ecoregion, 77d (Environmental Protection Agency, undated).
134. Subalpine fir-Engelmann spruce moist forest- In a small draw or gulch positioned within relatively low ridges of the inner Norther Cascades a lowland climax (or, perhaps, subclimax) forest had developed that was comprised of adult ("full-grown") Engelmann spruce, western hemlock, and mountain hemlock, and with younger trees of subalpine fir. The Very Tolerant subalpine fir (Wenger, 1984, p. 6) had the most apparent regeneration and, as was shown below, appeared to be the coniferous species that was ascending into dominance. Species composition was that of the climax subalpine fir-Engelmann spruce forest, but absence (or nearly so) of adult subalpine fir from local stands and instead presence of immature trees (large saplings to smaller poles) of this species was not typical of an old-growth climax forest of this cover (Eyre, 1980, p. 86). Western and mountain hemlocks were associate species in this forest tract. This was an exterior view of that mesic high montane or low subalpine conifer forest. It was near old-growth forest stage, if not old-growth, with sizeable volume of downed timber and almost no understorey except for conifer regeneration that was mostly subalpine fir.
The photograph presented here was taken from the berm of a road in order to show physiogonomy (to the extent possible) and general structure as apparent from an exterior perspective. There were natural openings (most appeared to be gaps or patches produced by death of one or more large trees) throughout this forest that had vegetation like that shown here. The forest vegetation of these gaps could not be photographed to the advantage of either detail or tree height as in this slide due to shorter distance from adjoining forest (ie. gaps were not long enough to get camera focal length for tree height and to eliminate shade cast by the nearby forest cnopy).
Some photographs from one such natural tree gap were presented farther below to show details of what understorey had developed in this lowland, mesic forest.
Regeneration at edge of forest consisted of all four major coniferous species, but regeneration was limited primarily to subalpine fir inside the forest under deep shade of thick canopy. It was not known why the Very Tolerant western hemlock (Wenger, 1984, p. 6) had much less regeneration than did subalpine fir, given that there were as much adult (mature) western hemlock as there were at the layer of forest canopy. The main shrub in forest vegetation shown in this photograph was common snowberry.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). There was no appropriate unit given by Brown et al. (1998, p. 36-37), but there should be an Engelmann Spruce-Alpine Fir Series under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) the same as there was for Rocky Mountain and Great Basin Subalpine Conifer Forest. North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
135. Forest on the edge- Edge of a climax Engelmann spruce-subalpine fir (western and mountain hemlocks, associate species) lowland forest showing regeneration of the climax dominants and a shrub layer dominated by common snowberry. Tree species present: mountain hemlock (left trunk), Engelmann spruce (largest trunk; slightly right of center), western hemlock (trunk to left an rear of the Engelmann spruce), and mountain hemlock (two side-by-side large trunks to right of the big Engelmann spruce). The shrub growing at base and up on butt portion of Engelmann spruce trunk was vine maple (Acer circinatum).
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). There was no appropriate unit given by Brown et al. (1998, p. 36-37), but there should be an Engelmann Spruce-Alpine Fir Series under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) the same as there was for Rocky Mountain and Great Basin Subalpine Conifer Forest. North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
136. Entrance to the forest- A natural walkway through boles of mature and larger young growth (poles) of a stand of Engelmann spruce and subalpine fir was presented in the first of these two slides. In this local stand Engelmann spruce and subalpine were co-dominant while western hemlock and mountain hemlock were present only as saplings at this imaginary entrance into a old-growth (or nearly) climax forest that developed in a small draw or gulch within a series of low ridges in the interior of the Northern Cascades. The second slide showed regeneration deeper inside this forest but in a small opening on the ground with a larger open space in the canopy. Common snowberry and Alaskla huckleberry grew around edges of this "hole" in the woods, but most vegetation was seedlings and saplings of coniferous species, especially the Very Tolerant subalpine fir which is the climax dominant in this forest cover type.
Looking in the "door" and down the forest "hallway" revealed presence of seedlings and saplings of both Engelmann spruce and subalpine fir, the current co-dominant species. Conifer regeneration was clearer and more obvious (shown at closer camera range) in the next set of two slides.
East Slope of Northern Cascades. Methow Valley Ranger District, Okanogan National Forest, Skagit County, Washington. June (third full day of summer). FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Engelmann Spruce-Subalpine Fir). There was no appropriate unit given by Brown et al. (1998, p. 36-37), but there should be an Engelmann Spruce-Alpine Fir Series under Cascade-Sierran Subalpine Conifer Forest biotic community (121.4) the same as there was for Rocky Mountain and Great Basin Subalpine Conifer Forest. North Cascades- North Cascades Highland Forests Ecoregion, 77b (Environmental Protection Agency, undated).
*Scrub Forests of the Coast Ranges
Scrub forests occurring along the Pacific Coast and outer edge of the Coast Ranges were treated under Pacific Northwest Forests.
*Pacific Douglas-fir Forest of the Coast Ranges
Douglas-fir forests (including industrial or commercial forests) of the Coast Ranges were treated under Pacific Northwest Forests.
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