Intermountain Forests

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The Intermountain Region or Intermountain West is the vast geographic portion of North America situated between the Sierra Nevada and Cascade Range to the west and the Rocky Mountains and associated mountain ranges to the east. In terms of physiographic provinces the Intermountain Region includes Basin and Range, Colorado Plateau, and Columbia Plateau. Given close proximity of forests in southeastern parts of the Basin and Range (eg. Mexican Highland and Sacramento Section) and in northeastern portions of the Columbia Plateau (eg. Blue Mountain and Harney Sections), these areas where conveniently--if physiographically imprecise--included with the Central and Southern Rocky Mountains and Northern Rocky Mountains, respectively.

Forests covered in this chapter were primarily those of the Great Basin and Colorado Plateau. Affinity of these forest with the Sierra Nevada-Cascade Range and the Rocky Mountains-Wyoming Basin was obvious floristically (range plant species and plant communities), climatically, geologically, and edaphically.

This treatment was consistent with that of the Intermountain Flora Project that produced the nearly epic-making Intermountain Flora. In defining the "floristic Intermountain Region" for their Intermountain West series Cronquist et al. (1972) explained that the selected geographic area as defined by the flora working was arbitrary and included political boundaries added atop natural floristic, geologic, climatic, etc. criteria. Specifically, the Intermountain West included the state of Utah "a priori" and drew the eastern boundary at the Utah-Colorado state line. Furthermore, the Intermountain Flora Project was defined as containing four distinct floristic divisions: the floristic Great Basin, Wasatch Mountains, Unita Mountains, and Colorado Plateau. The two large mountain ranges are of course part of the Middle (= Central) Rocky Mountains. Simply put, the North American Intermountain West can floristically and conveniently--if somewhat arbitrarily--be interpreted as the Great Basin portion of Basin and Range and the Colorado Plateau.

That was the format utilized in this chapter with treatment of range vegetation devoted (and limited) to natural forest communities and associated woodlands and shrublands. When these woody plant-dominated range plant communities were taken together the array of potential natural vegetation of mountains within the Intermountain Region could be viewed in context of Landscape Ecology. Such was the perspective in the following treatment of range cover types in Great Basin and Colorado Plateau montane forests.

Zonation of range plant communities in mountains of the Intermountain West is similar to the zonal vegetation in the Rocky Mountains and Sierra Nevada-Cascade Range. Consistent with coverage of woodland and forest range vegetation in these mountains some of the range types below were also included in other chapters like Pinyon-Juniper Woodland, Aspen, and Miscellaneous Scrub Types.

Most of the examples of various Intermountain Region range cover types presented below came from the Snake Range in Humboldt-Toiyabe National Forest much of which was transferred from the United States Department of Agriculture (Forest Service) to United States Department of Interior (National Park Serivce) to become Great Basin National Park.

1. Basin meets range- In ascending from the basin of Snake Valley up into the foothills of the Snake Range the range plant communities change from salt desert shrub dominated by the chenopod shrub, winterfat (Eurotia lanata= Ceratoides lanata), to a shrub steppe dominated by mountain big sagebrush (Artemisia tridentata ssp. vaseyana) and bunchgrasses of the Festucoideae or Pooideae and Eragrostoideae (fescue or bluegrass and lovegrass subfamilies). Adjecent to (and slightly higher in elevation than) the sagebrush shrub steppe is the pinyon-juniper woodland, the first (lowest elevation) range vegetation dominated by trees. On some range this change is abrupt, almost fenceline-like. On other ranges or different parts of the same range there is a gradual shifting change (ie. a progressive--and successive--transition) of range vegetation. An abrupt chane or discrete boundary was the condition shown here.

The diverse species composition of the two distinct range plant communities included the following major or important species for: 1) sgebrush shrubsteppe- mountain big sagebrush, rubber rabbitbrush (Chrysothamnus nauseosus), viscid or Douglas rabbitbrush (C. viscidiflorus), needle-and-thread (Stipa comata), galleta (Hilaria jamesii), Indian ricegrass (Oryzopsis hymenoides= Stipa hymenoides), slender wheatgrass (Agropyron trachycaulum), small cover of bluebunch wheatgrass (A. spicatum), cheatgrass or downy brome (Bromus tectorum), prickly poppy (Argemone munita), and plains prickly pear (Opuntia polycantha) and for 2) small conifer woodland- most but less of the preceding species plus the domiants singleleaf pinyon pine (Pinus monophylla) and Utah juniper (Juniperus osteosperma).

A good (and readily available) summary of the oneleaf pinyon pine-Utah juniper woodland is that of Vasek and Thorne in (Barbour and Major, 1995, ps. 810-814).

Great Basin National Park. White Pine County, Nevada. June, estival aspect. Elevation was approximately 7,000 feet. The sagebrush-shrubsteppe had the following designations: FRES No. 29 (Sagebrush Shrubland Ecosystem), K-50 (Wheatgrass-Needlegrass Shrubsteppe), SRM 402 (Mountain Big Sagebrush), Brown et al. (1998, p. 40) unit should be a Stipa Series 142.26, or if must use existing series, Ricegrass Series 142.23 of Great Basin Shrub-Grassland 142.2,and Artemisia tridentata ssp. vasyeana / Heterostipa comata Shrubland (Nevada Natural Heritage Program (26 September, 2003). Pinyon-juniper woodland had the following designations: FRES No. 35 (Pinyon-Juniper Shrubland Ecosystem), K-21 (Juniper-Pinyon Woodland), SRM 412 (Juniper-Pinyon Woodland), SAF 239 (Pinyon-Juniper), Brown et al (1998, p. 38) unit Pinyon-Juniper Series 122.71 of Great Basin Conifer Woodland 122.7, Pinus monophylla-Juniperus osteosperma / Artemisia tridentata Woodland (Nevada Natural Heritage Program, 26 September, 2003). Both range plant communities were in Central Basin and Range- Carbonate Woodland Zone Ecoregion,13q (Bryce et al., 2003).

2. A range plant "mixer"- Ecotone between shrub-bunchgrass steppe and pinyon-juniper woodland. There is often a narrow transition (= an ecotone) in plant -animal communities between the sagebrush shrubsteppe and the adjacent but ever-so-slightly-higher-in-elevation, small-conifer woodland. This mixed range plant community ("mixture" of species from shrub-bunchgrass steppe and pinyon-juniper woodland) is different in species composition, physiogonomy, and structure/function of either of the major zonal plant communities.In such a vegetational situation there is a diffuse boundary between adjoining range plant communities. Such ecotonal range vegetation was shown here.

Species included dominant singleleaf pinyon pine, Utah juniper, mountain big sagebrush, rubber rabbitbrush, viscid rabbitbrush, needle-and-thread, Indian ricegrass, galleta, slender wheatgrass, smaller cover of bluebunch wheatgrass, cheatgrass, prickly poppy, and plains prickly pear.

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation was about 7,000 feet. Great Basin National Park., White Pine County, Nevada. June, estival aspect. FRES No. 29 (Sagebrush Shrubland Ecosystem). K-50 (Wheatgrass-Needlegrass Shrubsteppe).SRM 402 (Mountain Big Sagebrush). Wheatgrss Series 142.21 Great Basin Shrub-Grassland 142.2 of Brown et al., 1998, p.40). Artemisia tridentata ssp. vaseyana / Psuedoroegneria spicata Shrubland (Nevada Natural Heritage Program, 26 September, 2003). Central Basin and Range- Carbonate Woodland Zone Ecoregion, 13q (Bryce et al., 2003).

Pinyon pine-juniper woodland is generally regarded as being in the Upper Sonoran Life Zone in the scheme of C. Hart Merriam (1890). The photographs of pinyon pine-Utah juniper-mountain sagebrush-perennial grass vegetation were at roughly 6,800 to 7,300 feet elevation.

3. Not large trees but a big invasion- Landscape-scale perspective (first slide) and close-in view (second slide) of successionally excessive recruitment of singleleaf pinyon and Utah juniper on a pinyon pine-juniper-mountain big sagebrush- perennial grass range community. This was a natural ecotone between mountain big sagebrush-needle-and-thread-galleta-Indian ricegrass shrubsteppe and a singleleaf pinyon pine-Utah juniper woodland that has the the preceding plant community as an undertstorey. Sadly, on thre range presented in these two photographs the tree component (layer) was increasing at unnaturally high rates and canopy cover at the expensive of both the shrub (big sagebrush, ruibber rabbitbrush , and viscid rabbitbrush) and native perennial grass layers. This conifer invasion was likely due in most immediate time frame to cessation of fire (a consequence of relatively successful "total fire suppression" by whhite man. The resultant range plant community was a departure from potential natural or climax vegetation and was clearly range degradation by the process of plant retrogression. The Eurasian annual, Cheatgrass, was also present but not to the cover or density to cause major concern. Forbs were scarce.

Small conifers in foreground of both photographs were the nut or fox pine, singleleaf pinyon pine. Both conifer species are native plants and comprise the natural dominant layer of this potential lower foothill-montane coniferous woodland. Problem is "too much of a good thing" of these two species and conmittant decline in relative proportions of composite shrubs and perennial grasses.

The National Park Service should initiate and carry out an aggressive prescribed fire program (should have done so years ago). Otherwise this woodland-shrub-grass range vegetation will become a conifer forest which is a drastic departure from climax vegetation (and, ultimately, from the animal portion of this natural range ecosystem). On this rangeland the real friends of Smokey Bear do play (work) with matches or, better yet, drip torches.

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation approximately 7,000 feet. Various ecosystem, potential natural vegetation, SRM and SAF cover types, ecoregion, etc. were cited in caption 2.

4. Proof of conifer invasion- Young singleleaf pinyon pine growing in a mature mountain big sagebrush where the large seed of this stone pine fell and was rolled by gravity, blow by wind, or carried by some animal.

Great Basin National Park, White Pine County, Nevada. June, estival aspect.

CAUTION: Great care in proper use of prescribed burning will be essential on this, and similar, pinyon-juniper-sagebrush-grass range. If fire is too intense (burning is done at the wrong season or under under improper weather conditions) excessive loss of sagebrush or even grass could occur. A simultaneous disastrous outcome could be increase in cheatgrass.

Nonetheless, there is no practical alternative to prescribed fire as tool for range improvement (restoration of natural range vegetation). And prescribed burning should be done as soon as possible. As conifers like the one shown here become larger there will be more fuel and a hotter fire which can only increase likeli hood of excessive damage to shrubs and/or native grasses.

5. Excessive conifer recruitment was still a problem- A different singleleaf pinyon pine-Utah juniper woodland community was the one shown here on a xeric, steep, predominant west slope. This natural range vegetation is the same as the one described above and at SRM-SAF range types, FRES ecosystem, Kuchler "vegetation type", etc. cited in caption 2 except this is Pinus monophylla-Juniperous osteosperma / Sparse Understory Woodland (Nevada Natural Heritage Program, 26 September, 2003). This climax vegetation naturally lacks as many shrubs and herbaceous plants in lower layers. Only deeper-rooted trees and a few "lucky" (or "tough") understorey plants can persist on this range site characterized by shallow soil, steep slopes, and low soil moisture. Once again, however, excessively high rates of conifer recruitment were occurrring so that in the not-too-distant future the crown cover of conifers will be excessive and effective in preventing limited precipitation from penetrating and percolating into the shallow soil of this site. The few shrubs and grasses will become fewer as time goes by. (Again, this development is due largely to absence of natural fire.)

Loss of sagebrush shrubsteppe by excessive invasion of Utah junifer and singleleaf pinyon pine has been (and of this writing continues to be) a ecosystem-threatening development on this public range.This was an example of afforestation due to human inaction. It was improper management of the range ecosystem owned by all citizens and managed by public servants who "hold this public in trust". Afforestation is "the establishment of a forest or stand in an area where the preceding vegetatoion or land use was not forest" (Helms, 1998).

BURN IT OR LOOSE IT! Do not forget to heed the above caution.

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation was roughly 7,000 feet (6,800 to 7, 200 feet).

6.Brush-infested Great Basin pinyon-juniper woodland raange- General view of a degraded range plant community consisting mostly of overstocked Utah juniper and big sagebrush with widely scattered remnants of Indian ricegrass. The potential natural vegetation was Utah juniper-big sagebrush-Indian ricegrass woodland or savnna. This was designated as Juniperus ostersperma / Artemisia tridentata / Achnatherum huyemnoides Woodland by National Vegetation Clsssification for Nevada (Nevada Natural Heritage Prograqm , 26 September, 2003).

In absence of natural (prehistoric) fire regime Utah juniper was overstocked and big sagebrush had increased to excessive canopy cover resulting in loss of almost all of the herbaceous understorey of Indian ricegrass, galleta, slender wheatgrass, and western wheatgrass. Past overgrazing can--more times than not--be listed as a factor in loss of herbaceous cover, but in this example overgrazing was not a factor as will be proven conclusively in slides that follow. Rather the single determining problem was unnatural excess cover of Utah juniper. The cause of range deterioration in this plant community was overstocking of juniper. No "ifs, ands, or, buts about it".

Overstocking and overstocked are standard terms used in Forest Science and the profession of Forestry. Forestry Terminology (Munns, 1950) published by Society of American Foresters (SAF) provided this definition: "Overstocked. A condition of stand or forest indicating more than normal or full stocking would require. Domints in such stands have narrow rings indicating suppression." The subsequent and expanced edition of this dictionary was published jointly by the SAF and Food and Agriculture Organization of the United Nations (Ford-Robertson, 1971) gave these definitions: "Stocking (2) (silviculture/ management) In a forest, a more or less subjective indication of the number of trees as compared to the desirable number for best results e.g. overstocking ...(3) More precisely, a measure of the propoortion of the area actually occupied by trees, expressed e.g. in terms of stocked quadrants or % canopy closure, as distinct from their stand density..."

Put in terms appropriate for the lesson presented here: there were too many trees and their crowns covered too much of the soil surface to be natural, climax, potential natural, (or whatever) vegetation. While overgrazing, commercial activity, transportation, etc. may have been factors that enhanced juniper establishment, ultimately it was absence of fire that was responsible for the on-going overstocking and, subsequent, shading out of native perennial grasses. That was proven in slides below. First it was established that there was overstocking of conifers (ie. too many trees and too much canopy cover). This was shown even more dramatically in the next photograph.

Note continuing recruitment of Utah juniper by presence of a seedling size tree in left foreground (left margin). Note also some remnant grasses (mostly Indian ricegrass) visible as tan-colored clumps. Relatively high cover of big sagebrush was also evident, but that was not the main cover component that restricted growth and reproduction of grasses and other herbaceous species.

Tintic Valley, Juab County, Utah. June. FRES No. 35 (Pinyon-Juniper Shrubland Ecosystem), K-21 (Juniper-Pinyon Woodland), SRM 412 (Juniper-Pinyon Woodland), SAF 239 (Pinyon-Juniper), Brown et al (1998, p. 38) unit Pinyon-Juniper Series 122.71 of Great Basin Conifer Woodland 122.7, Juniperus osteosperma / Artemisia tridentata / Achnatherum hymenoides Woodland (Nevada Natural Heritage Program, 26 September, 2003). Central Basin and Range- Sagebrush Basins and Slopes Ecoregion, 13c (Woods et al., 2001).

7. What happens when Smokey's friends do not "play with matches" (or drip torches or, at least, let Mother Nature do so)- Two views of interior of an overstocked Utah juniper woodland with very little shrub or herbaceous layers. Excessive and unnatural crown cover of juniper excluded native shrubs (including the natural dominant, big sagebrush) and grasses (especially Indian ricegrass) from what should have the natural vegetation of a Utah juniper-big sagebrush-Indian ricegrass woodland or savanna. Instead this degraded woodland range had become through mismanagement (namely fire exclusion/fire suppression) a single-species stand of Utah cedar. This was if not by design at least by default a monoculture just as surely as if the junipers had been planted like an orchard. This vegetation had become a one species forest instead of a tree-shrub-grass woodland or savanna.

A few struggling plants of Indian ricegrass were visible in foreground of first of these two photographs. One such plant of Indian ricegrass was shown in the second photograph. Absence of big sagebrush in the stand of juniper should be observed. Compare this group of plants to that in the immediately preceding slide and observe that big sagebrush was present only outside (up to edge) of the juniper forest.

Smokey Bear was here: exactly where we did not need the misguided bruin.

Tintic Valley, Juab County, Utah. June, early estival aspect. FRES No. 35 (Pinyon-Juniper Shrubland Ecosystem), K-21 (Juniper-Pinyon Woodland), SRM 412 (Juniper-Pinyon Woodland), SAF 239 (Pinyon-Juniper), Brown et al (1998, p. 38) unit Pinyon-Juniper Series 122.71 of Great Basin Conifer Woodland 122.7, Juniperus osteosperma / Artemisia tridentata / Achnatherum hymenoides Woodland (Nevada Natural Heritage Program, 26 September, 2003). Central Basin and Range- Sagebrush Basins and Slopes Ecoregion, 13c (Woods et al., 2001).

8. What happens when Smokey's friends do "play with matches"- Natural "fireline" contrast: seven years (six growing seasons) before this photograph a wild fire (Tintic Valley Railroad Fire of 1999) burned this degraded, bare understorey, overstocked Utah juniper woodland. Source of ignition of this fire was thought to be sparks from railroad truck grinders, hence known as the Railroad Fire of 1999 (Thompson, 2002). Wild fire burned up to edge of this stand of Utah juniper and stopped at the location shown in this slide. The burnt-over juniper woodland range to the right served as a natural treatment while the unburnt portion on the left served as a de facto "control plot".

What a difference a good fire makes, and even a wild fire can be good! Results of this unplanned burn (1999 Railroad Fire in Tintic Valley) were dramatic Outcome was mostly dead juniper trees and release of native grasses such as the dominant Indian ricegrass that was growing so beautifully in right foreground (beneath beautifully dead Utah juniper). Utah juniper does not resprout. If a fire is hot enough to kill the crown of a Utah uniper it will "stay dead".

Important point: all range revetation was by secondary plant succession. This was private property that was not treated by reseeding following the 1999 Railroad Fire.

9. Good-lookin' Utah juniper and even better-lookin' grass or seven years after the 1999 Railroad Fire in Tintic Valley Utah- Three photographs taken seven years (six growing seasons) after a wild fire burned an overstocked Utah juniper-big sagebrush-Indian ricegrass woodland or savanna from which native shrub and perennial grass layers had been excluded by excessive crown (= canopy) cover of juniper. After ten growing seasons native grasses had made a remarkable recovery and big sagebrush was becoming re-established. These three photographs of a wild fire "treatment" were taken about 200 yards and less than a quarter-hour from the "control plots" (the part of this same juniper-sagebrush-grass woodland range that was unburned).

It should be noted that some Utah juniper were somehow spared from the fire even inside large burnt portions. These adult trees served as a seed source for reinvasion of juniper on this range site. This will result in eventual development back into a woodland with restored woodland physiogonomy, structure, and function. This was an example of Clements "dynamic plant ecology". In other words, maintenance of the juniper-shrub-grass woodland will require maintenance fires (or some other substitute like chain saws for instance).

Most of the dramatic increase in grass cover had been that of the climax dominant, Indian ricegrass, but galleta, slender wheatgrass, and western wheatgrass had also increased to almost unbelieveable levels of biomass and foliar cover. Cheatgrass was not a common component of this post-fire range plant community though, of course, cheatgrass is usually present on Great Basin range (unless cedars shade it out). Desert crested wheatgrass was also present on two sides of the border of this range that were adjacent to a state highway.

The main forb on this burnt-over range was the native, perennial composite fewflower wirelettuce (Stephanomeria pauciflora). Wirelettuce was a major herbaceous species at local scale. Large clumps of this species were readily distinguishable in the first of these three slides (one in extreme left foreground).

NOTE: This rangeland was not treated by post-fire rehabilitation. All recovery of range vegetation was natural. This was private property. This range did not receive post-fire reseeding treatments as were applied on range managed by Bureau of Land Management that were also burned by the 1999 Railroad Fire. Readers can find discussion of those rehabilitation treatments in Thompson (2002).

From comparisons across all photographs--seven years (six growing seasons) post-burn--it was readily apparent that area of bare ground was higher under unburned junipers than on burned range.

6. Where did the shrubs go? (Ah, but look at all the great grass.)- Foreground range vegetation was Utah juniper woodland with a dense understorey of grass and no shrub layer. Dominant grass was the cespitose needle-and-thread. There was also good representation by galleta, Indian ricegrass, and, as always some cheatgrass or downy brome. The shrub layer in background and around crown drip line of juniper was dwarf mountain-mahogany (Cercocarpus intricatus). It was not known if absence of shrubs in foreground was due to fire, unsuitable habitat (eg. soil), competition with grass, or some other phenomena. Pinyon pine was absent from this range vegetation.

This range plant community was partially described by Western Heritage Task Force (Bourgeron et al., 29 August, 1994) and Nevada Natural Heritage Program (26 September, 2003) as Juniperus osteosperma / Cercocarpus intricatus Woodland. Exclusion of the herbaceous layer comprised of native grasses (and very little cheatgrass) rendered such unit of classification incomplete. FRES No. 35 (Pinyon-Juniper Shrubland Ecosystem). K-21 (Juniper-Pinyon Woodland). Variant of SRM and SAF 412 (Juniper-Pinyon Woodland). Variant of SAF 239 (Pinyon-Juniper). Brown et al. (1998, p. 38) Pinyon -Juniper Series 122.71 of Great Basin Conifer Woodland 122.7. Central Basin and Range- Carbonate Woodland Zone Ecoregion, 13q (Bryce et al., 2003).

Beaver County, Utah. June, estival aspect.

7. Co-dominants of Great Basin conifer woodland- Singleleaf pinyon pine (Pinus monophylla), left and largest tree, and Utah juniper (Juniperus osteosperma), other trees. Also present were mountain big sagebrush, needle-and-thread and Indian ricegrass.

Great Basin National Park, White Pine County, Nevada. June.

8. Picturesque specimen- Singleleaf pinyon pine on a ecotone of pinyon pine-juniper woodland and mountain big sagebrush- mixed grass shrubsteppe. This range vegetation was shown above.

Delightful natural history of the pinyon pine is that by Lanner (1981; 1983, ps.30-34). Also for fireside reading was the interesting account by Peattie (1959, p.s 73-75). Peattie also took a turn on Utah juniper (1959, ps. 263-266) as did Lanner (1983, ps.112-115).

Great Basin National Park, White Pine County, Nevada. June.

9. Tweedle Tree One and Tweedle Tree Two- Two trees of Utah juniper (Juniperus osteosperma). Other range plants included big sagebrush, Great Basin wildrye (Elymus cinereus), Indian ricegrass, and viscid rabbitbrush.

Tooele County, County, Utah. June.

10. Pinyon pine, nut pine, or Colorado pinyon (Pinus edulis)- The "squatty" form (general morphological appearance) of the classic pinyon pine species. This is the one most feasted on first by American Indians and, later also, the white man. The "nuts" (not nuts, of course, but the naked, stone seeds) of P. edulis are still gathered, then roasted and eaten with considerable enjoyment. Seeds of this and other "nut" or "stone" pines are important concentrates for wildlife and even livestock, especially sheep.

This specimen was growing on a shamefully overgrazed range in Las Animas County, Colorado. August. Overgrazing--by cattle in this instance--facilitated the dispersion (including seed movement by surface water as excessive runoff or overland flow), germination/emergence, and reduced competition with more palatable mid- and shortgrass species.

11. Staminate cones and needles of singleleaf pinyon pine- Characteristic male cones were presented with the unique one needle per fascicle in singleleaf pinyon pine.

Great Basin National Park, White Pine County, Nevada. June, anthesis in full-swing.

12. Immature pistillate cone of singleleaf pinyon pine- Female woody, seed-bearing structure (= cone) at a state of immaturity. Singleleaf pinyon pine is the pine species that is co-dominant in much of the pinyon-juniper Great Basin woodland. The field- identifying characteristic of one needle per fascicle was obvious.

Great Basin National Park, White Pine County, Nevada. June.

13. Empty woody nest- Female cone of singleleaf pinyon pine from which seeds had been shed. The pinyon pines (there are four species in the pinyon-juniper woodland of the Western Range) are in the Pinus group known as nut pines. Some students include the nut pines in a generic group called stone pines. Reference to stone is in regard to the large seeds, several of which are edible and highly prized by numerous animals including man. Obviously the naked seeds of gymnosperms are not nuts, which are a kind of woody fruit of angiosperms, but the use of "nut" gets across the idea or mental picture of large, hard seeds. Bearing of such large seeds contrast shrapely with the usual parchment-like scale seeds of other pines. The fact that such seeds are favored food by different animals helps assure sexual reproduction of the nut pines. (As shown above this reproduction is sometimes too successful in absence of phenomenon or factors that regulate pine populations to their proper stocking rate.)

Presence of only one needle per fascicle in singleleaf pinyon pine was also shown in this photograph.

The pinyons or nut pines (in more restricted usage) are included in the soft or wihte pine group (subgenus Haploxylon).

Great Basin National Park, White Pine County, Nevada. June.

14. Single-leaf pinyon pine (Pinus monophylla)- Single-leaf or one-leaf pinyon is a widely distributed "nut" or "stone" pine with a biological range from northern Mexico to southern Idaho. It is especially "happy" to reside over much of southern California from the east-side pine type throughout both the Mojave and Sonoran Desert Regions.

The individual single-leaf pinyon preented here was growing in Riverside County, California along the famed Palms to Pines Drive, California State Route 74. What a wonderful route along which to study the Life Zones of C.Hart Merriam and restore one's soul. Highly recommended and happy motoring. June.

15. Still in the Upper Sonoran Life Zone- At elevations immediately above the pinyon pine-juniper woodland or, as the upper elevational limit of the pinyon-juniper woodland, there is frequently (as shown here in the Snake Range) a unique kind of range vegetation consisting of juniper, with or without pinyon pine, and low sagebrush (Artemisia arbuscula) as a shrub or low woody layer plus an herbaceous layer of various species (especially grasses). Two examples of this grass-low shrub-pinyon juniper woodland were shown in these two photographs. The sample of this variant of Great Basin conifer woodland lacked pinyon pine as singleleaf nut pine had "fallen out way back down the mountain". So this was a Utah juniper-low sagebrush woodland.

Other shrubs included a few "lingering" individuals of mountain big sagebrush (one was prominent in center foreground of second slide) and various other low shrubs including "runt"-looking viscid or Douglas rabbitbrush, broom snakeweed (Gutierrezia sarothrae), and hairy golden aster (Heterotheca villosa). There were a few individuals of curl-leaf mountain -mahogany (Cercocarpus ledifolius) which was the dominant plant of the elevation-based zonal vegetation immediately above this range plant community. Some mountain-mhogany plants were visible in second photograph at left margin by juniper. Composite forbs included the complicated flebane crowd (Erigeron spp.) and basin butterweed or Unita groundsel (Senecio multilobatus). Dominant grass was the sparsely populated muttongrass, mutton bluegrass, or Fendler's bluegrass (Poa fendleriana). Needle-and-thread, Indian ricegrass, and bluebunch wheatgrass were limited to moist microsites (often near junipers).

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation was in approximate range of 7,800 to 8,300 feet. FRES No. 35 (Pinyon-Juniper Shrubland Ecosystem) K- 21 (Juniper-Pinyon Woodland). SRM 412 (Juniper-Pinyon Woodland). SAF 239 (Pinyon-Juniper). Brown et al. (1998, p. 38) Pinyon-Juniper Series 122.71 of Great Basin Conifer Woodland 122.7. Juniperus osteosperma / Artemisia arbuscula Woodland (Nevada Natural Heritage Program, 26 September, 2003). Central Basin and Range- Carbonate Woodland Zone Ecoregion, 13q (Bryce et al., 2003).

16. Upper elevation of Upper Sonoran Life Zone- Curl-leaf mountain-mahogany shrubland was the range vegetation at the highest elevation of the Upper Sonoran Life Zone in the Snake Mountains. These two landscape-scale photographs depicted characteristic curl-leaf mountain mahogany scrubland in the Snake Range relative to zones (= belts) of range plant communities in these mountains. Zonal range vegetation was determined to large extent by factors (precipitation, temperture, soils, degree of slope, winds, etc. ) associated with differences in elevation and, to lesser degree, direction of slope (eg. north, south, west slopes).

Distant range vegetation that was dark green in color was tree-dominated range plant communities. That which was below (lower in elevation) this mountain-mahogany belt were various communities dominated by singleleaf pinyon pine and juniper (primarily Utah juniper). Natural plant communities above this zone of mountain mahogany scrub (higher in elevation) were vrious forest cover types (ponderosa pine-Douglas-fir, quaking aspen, Englemann spruce, limber pine, and bristle cone pine). The bare area on the distant mountain, which was Wheeler Peak, was the alpine range ecosystem above timberline.

Some of the curl-leaf mountain-mahogany plants were of heights (as tall as 15-25 feet) more typical of trees than shrubs, but almost all such plants were of the multi-stemed habit characteristic of shrubs. This woody plant-dominated (based on canopy cover, physiogonomy, plant height) vegetation was therefore defined precisely as shrubland or scrub and not woodland.

There was one-- sometimes two-- layer(s) lower layers of plants. Shrubs that were relative common in lower woody layer(s) included low sagebrush (which was the understorey dominant in the adjacent and lower Utah juniper-low sagebrush range community), viscid rabbitbrush, hairy golden aster, and fringed sagebrush (Artemisia frigida). Composites were clearly "in control" of lower shrub layer(s).

The herbaceous layer of this range vegetation was dominated by muttongrass of Fendler's bluegrass. The associate herb was the range forb, lanceleaf phacelia (Phacelia hastata= P. leucophylla). Many of the same range forb species grew on this belt of mountain-mahogany scrub range and the adjacent zone of Utah juniper-low sagebrush-grass range.

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation was roughly 8,000 to 8,500 feet up to as high as 9,000 feet on dry south and west slopes. FRES No. 34 (Chaparral-Mountain Shrub Shrubland Ecosystem). K- 31 (Mountain-mahogany-Oak Scrub), mountain-mahogany variant thereof. SRM 415 (Curlleaf Mountain Mahogany). Mountain-Mahogany Series 132.12 of Great Basin Montane Scrub 132.1 of Brown et al. (1998, p. 39). Cercocarpus ledifolius Shrubland Alliance is as far as can go because National Vegetation Classificatioin for Nevada (Nevada Natural Heritage Program, 26 September, 2003) did list a community (association) of this composition. Central Basin and Range- Carbonate Woodland Zone Ecoregion, 13q (Bryce et al., 2003).

17. Scrub range in the Snake Range- Interior of the curl-leaf mountain-mahogany-low shrub-muttongrass scrubland range presented in the two immediately preceding photographs. The multi-stemmed habit of mountain-mahogany plants was pronounced. Herbaceous and woody layers in the understorey of this range vegetation varied greatly at local (microclimate) scale. Single species of plants tended to form local populations or colonies (Clements, process of aggregation or "birds of a feather flock together"). This was particularly so for lanceleaf phacelia and muttongrass. Perhaps this was a combination of plant dispersal characteristics and considerable vriation in edaphic habitat at scale of microsites.

Several plants of fringed sage were noticable in foreground of both photographs.

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation was approximately 8,000 to 8,500 feet up to 9,000 feet on south and othr dry slopes. FRES No. 24 (Chaparral-Mountain Shrub Shrubland Ecosystem). K-31 (Mountain-mahogany-Oak Scrub), "pure" mountain-mahogany variant thereof. SRM 415 (Curlleaf Mountain Mahogany). Mountain-Mahogany Series 132.12 of Great Basin Montane Scrub 132.1 of Brown et al. (1998, p. 39). Cercocarpus ledifolius Shrubland Alliance is "it" because there was not an association level in National Vegetation Classification for Nevada (Nevada Natural Heritage Program, 26 September, 2003) having this composition. Central Basin and Range- Carbonate Woodland Zone Ecoregion, 13q (Bryce et al., 2003).

18. Curlleaf mountain-mahogany (Cercocarpus ledifolius)- Typical plant of this shrub species in the Snake Range of the Great Basin. Description of this species as to browse value was in the Range Plant Handbook (Forest Service, 1940, B50) as well as Dayton, 1931, p. 45) and Sampson and Jespersen (1963, ps. 80-81). See also Lanner (1984, ps.180-182) for a brief natural history.

Snake Range, Great Basin National Park, White Pine County, Nevada. June, pre-bloom stage..

19. Foothill neighbor- Another plant of cutleaf mountain-mahogany. This one was a neighbor to a population of Gambel's oak (Quercus gambelii) and a few New Mexico locusts (Robinia neomexicana) on a seep in the foothills of the Front Range of Southern Rocky Mountains.

Huerfano County, Colorado. Mid-June (late spring), pre-bloom stage.

20. Curlleaf mountain-mahogany - Typical columnar habit of curlleaf mountain mahogany. According to Dayton (1931, p. 45) this is probably the largest and most treelike of the Cercocarpus species. In addition, it has evergreen leaves which increase it's browse value during winter months.

Rock Canyon, Utah County, Utah. June.

21. Leaders of curlleaf mountain mahogany- Bark and leaves of curlleaf mountain mahogany. Rock Canyon, Utah County, Utah. June.

C. montanus and C. ledifolius were described in the various editions of North American Range Plants (Stubbendieck et al., 1981, 1982, 1986, 1992) and the Range Plant Handbook (Forest Service, 1940, B49-B51) as well as miscellaneous state and regional references on browse plants such as for California (McMinn, 1939, ps. 205-209; Sampson and Jesperson, 1963, p. 78-81) and eastern Oregon and Washington (Hayes and Garrison 1960, ps.178-180).

22. Muttongrass, Fendler's bluegrass, or mutton bluegrass (Poa fendeleriana)- This delightful festucoid grass was the dominant herbaceous species on a curl-leaf mountain-mahogany scrubland range.Muttongrass (descriptive common name for value of this species as sheep forage) is one of the most important of all bluegrasses on the Western Range. A valuable practical description of muttongrass was provided by the Range Plant Handbook (Forest Service, 1940, G100). This remarkable species ranges from the Chisos Mountains of Trans Pecos Texas to British Columbia.

Muttongrass has deservedly been on the Society for Range Management "200 list" for the Intercollegiate Range Plant Contest.

These fine specimens were in the understorey of a curl-leaf mountain-mahogany scrub range (on which it was the dominant herbaceous species). Snake Mountains, Great Basin National Park, White Pine County, Nevada. June, grain-ripe stage.

23. Panicles of muttongrass- Panicles with large spikelets of full of ripe grain in Poa fendeleriana. Spikelets of muttongrass are some of the largest of any bluegrass species in North America. Fendler's bluegrass is a major range forage species in the mountains of the Great Basin.

Snake Range, Great Basin National Park, White Pine County, Nevada. June, "seed-ripe" phenological stage

24. Lanceleaf phacelia (Phacelia hastata= P. leucophylla)- This was the most abundant forb on the curl-leaf mountain-mahogany scrubland range featured above. This forb species was also locally common on the Utah juniper-low sagebrush range that was adjacent to but slightly lower in elevation than the mountain-mahogany scrub zone.

Incidentially, Phacelia is one of the more numerous genera of range forbs in the Intermountain Region. Welsh et al (1993) described 39 species of Phacelia in Utah, though not all of them occurred in or were limited to the Great Basin.

Snake Range, Great Basin National Park, White Pine County, Nevada, June, full-bloom phenology.

25. Lanceleaf stonecrop (Sedum lanceolatum)- This showy little fellow was growing on just one more rockpile on the curl-leaf mountain mahogany scrub range shown above. This species was also found on the Utah juniper-low sagebrush range just below the mountain-mahogany zone.

The formerly obscure Crassulaceae became well-known (at least indirectly) to students of Plant Physiology when the third photosynthetic pathway was discovered in members of this family with the metabolic pathway being named Crassulacean Acid Metabolism. Ain't range plants groovy!

Snake Range, Great Basin National Park, White Pine County, Nevada. June, "in all her glory" (ie. full-bloom stage).

26. Basin butterweed or Unita groundsel (Senecio multilobatus)- Senecio species are some of the major range composites of mountain grazinglands throughout western North America. Identification of Senecio species can be challenging, but this is one of the more widespread and common ones. Basin butterweed grew on both the Utah juniper-low sagebrush and curl-leaf mountain-mahogany scrub range zones in the Snake Mountains.

The large fuzzy leaves in right background belonged to the introduced, now naturalized, and widely distributed velvet mullein (Verbascum thapsus).

Great Basin National Park, White Pine, Nevada. June, full-bloom stage of phenological development.

27. Inflorescences of basin butterweed or Unita groundsel- Flowers and fruit (achene) on the individual shown in the preceding photograph. Great Basin National Park, White Pine County, Nevada. June.

28. Scarlet gilia or sky rocket (Gilia aggregata= Ipomopsis aggretata)- This member of the Polemoniaceae (Phlox family) grew on both the Utah juniper-low sagebrush and curl-leaf mountain-mahogany range plant communities. This was "no big deal" beacause according to Welsh et al. (1993, p. 519) this species "... is practically ubiquitous in Utah". So sky rocket is bound to be common across the line in the neighboring Silver State.

Welsh et al. (1993, p. 519) also remarked that scarlet gilia is a "strikingly beautiful species, with the odor of a skunk.." If that combination of features does not merit a place with neighboring range forbs what would it take? Besides, Hermann (1966, ps. 231-232) included this species (complete with figure) in Notes on Western Range Forbs so by golly it's here too.

29.Trees in the Transition Zone- In most mountains of the Western Range Region (from the Rocky Mountains to the Pacific Slope) ponderosa pine or western yellow pine (Pionus ponderosa) forms zonal forests that typically have well-developed understories. This western yellow or ponderosa pner belt usually develops above the pinyon-juniper or associated woodlands and/or shrublands. Sampson (in Tridestrom, 1925, ps. 28-29) described this as the "yellow pine and oak brush belt" for the mountains of Utah and Nevada identifying this belt as the "transition zone". Coniferous forests dominated by ponderosa pine and associated trees, notably Douglas-fir (Pseudostuga menziesii), typically form the lowest elevation forests in the mountains of western North America. This is classically the Transition Life Zone of Merriam (1890). Inclusion of ponderosa pine forests in the Tranzition Life Zone is traditional (recall again this usage by Sampson cited earlier).

In the Snake Range the ponderosa pine-Douglas-fir transition zone forest developed as the next zonal range plant community above the curl-leaf mountain-mahogany belt. Mountain-mahogany was therefore a key component and indicator woody species in this lowest-elevation forest in the Snake Range (three or four shrubs of curl-leaf mountain-mahogany were conspicuous at the lowest edge of trees in the second of these two photographs). The low-growing, mat-like shrub with bright green leaves in foreground of both slides was greenleaf manzanita (Arctostaphylos patula). This species was "for all intents and purposes" the only shrub growing on the rockiest of sites and openings among the forest (foreground of both photographs).

The first of these slides was a wide-angle view of a stand of mature ponderosa pine with both young trees of both pine and Douglas-fir. Ponderosa pine is one of the most fire-tolerant of western conifers. It seemed likely that establshment (invasion) of Douglas-fir in the understorey of the ponderosa pine stand had been more plentiful under the existing policy (direct and de facto) fire suppression. This forest had been under management by the Forest Service until 20 years before this photograph was taken. A policy of fire suppression had been in effect for three-quarters of century up until two decades ago when the "let burn" polikcy of the National Park Service supplanted the standard fire management practice of "total suppression". Douglas-fir are fairly fire tolerant if they become established long enough to reach ages approaching maturity when their thick bark provides substantial protection from surface (not crown) fires.

A number of Douglas-fir trees with meaningful crown cover were present in the forest stand shown in the second slide. Tree along entire left margin of first slide was Douglas-fir. There were also some white fir (Abies concolor) farther back in the interior of forest vegetation shown in both slides.White fir is also much less tolerant of fire than is ponderosa pine. There were also even fewer limber pine (Pinus flexilis), a dominant species of forest communities at higher elevations here in Snake Range.

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation was about 8,500 to 9,000 feet and extending up to 9,500 feet on drier sites (eg. south slopes). FRES No. 21 (Ponderosa Pine Forest Ecossystem). K-10 (Western Ponderosa Foresst). No SRM rangeland cover type for ponderosa pine in Great Basin. SAF 237 (Interior Ponderosa Pine).In Brown et al. (1998, p. 37) would probably be Pinus ponderosa-Abies concolor Association 122.626 or Pinus ponderosa-Pinus flexilis Association 122.627 of Yellow Pine Series 122.62 under Rocky Mountain Montane Conifer Forest 122.6 becuse these author previously aligned higher-elevation conifer forests of Great Basin with Rocky Mountains. Pinus ponderosa Woodland Alliance and no farther in Nevada Natural Heritage Program (26 September, 2003), but Pinus ponderosa / Arctostaphylos patula for Utah (Bourgeron et al., 29 August, 1994). Central Basin and Range- Carbonate Woodland Zone Ecoregion, 13q (Bryce et al., 2003).

30. Three woody layers on a pile of rocks- Two "photo-plots" of a ponderosa pine-curl-leaf mountain-mahogany-greenleaf manzanita forest range. Closer views of rorest range vegetation shown in two immediately preceding photographs. (Note the same small curl-leaf mountain-mahogany in first of these slides as second of the two above slides.) Geenleaf manzanita was dominant shrub, but lower of two shrub layers (the otherand taller shrub layer formed by mountain-mahogany).

Ponderosa pine dominant at higher age classes, but Douglas-fir was more common locally at younger age classes. Some white fir and "lost" limber pines from higher-elevation forest communities.

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation was about 8,500 to 9,000 and up to 9,500 feet on xeric sites (eg. south and west slopes, shallow soils). FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-10 (Western Ponderosa Forest). No appropriate SRM rangeland cover type. SAF 237 (Interior Ponderosa Pine). In Brown et al. (1998, p. 37) would most likely be Pinus ponderosa -Pinus flexilis Association 122.627 or Pinus ponderosa-Abies concolor Association 122.626 of Yellow Pine Series 122.62 in Rocky Mountain Montane Conifer Forest 122.6 because higher-elevation conifer forests of Great Basin were aligned with Rocky Mountains. Pinus ponderosa Woodland Alliance is as far as goes (Nevada Natural Heritage Program, 26 September, 2003), but Pinus ponderosa / Arctostaphylos patula (Bourgeron et al., 29 August, 1994). Central Basin and Range- Carbonate Woodland Zone Ecoregion 13q (Bryce et al., 2003).

31. Layered Great Basin ponderosa pine forest- Close-in view of a local plant assemblage of the western pine forest that was presented in the first photograph that introduced this ponderosa pine forest range (ie. four slides back). Three very old ponderosa pine patriarchs are above two single stems of curl-leaf mountain-mahogany and dense greenleaf manzanita. Three woody speceis; three layers of vegetation

Snake Range, Great Basin NationalPark, White PIne County, Nevada. June, estival aspect. Elevation for range vegetation of this species composition was about 8,500 to 9,000 feet. This Ecosystem, Kuchler "vegetation type", SAF forest cover type, association designations, etc. were cited in the two preceding captions.

32. Great Basin Douglas-fir- Local stand of interior Douglas-fir that had established on a climax ponderosa pine-curl-leaf mountain-mahogany-greenleaf manzanita forest range. In this same tract of forest there werealso a few white fir and limber pine, two species that were common in forest communities higher up here in the Snake Range.

It was not know if a mixture of Douglas-fir and ponderosa pine constituted potential natural vegetation or, alternatively, if the less fire tolerant Douglas-fir had invaded in absence of natural fire much as the pinyon pine and Utah juniper were doing below. When this forest was under management by Forest Service as Humboldt-Toiyabe National Forest (for three-quarters of a century up until 20 years before this photograph) the prevailing fire policy was one of "total fire suppression". It was likely that many, if not most, of the Douglas-fir, white fir, and few limber pine in this Transition ponderosa pine forest had been able to invade from the Canadian Life Zone (the next Merriam zone above the Transition) in absence of regular surface fires. In other words, cessation of forest fire permitted the associate species, Douglas-fir, and two minor conifer species to "move down the mountain slope" into somewhat drier habitat(s) because this environment had become less hostile with elimination of a major controlling climatic factor.

Great Basin National Park, White Pine County, Nevada. June, estival aspect. Elevation for this forest range community in Snake Range varied around 8,500 to 9,000 feet or so. Designations as to ecosystem, Kuchler "vegetation type", SAF forest cover type, associations, etc.were cited in the first two captions for Great Basin ponderosa pine forest. This local stand was a Pseudotsuga menziesii / Arctostaphylos patula association (Bourgeron et al., 29 August, 2003).

33. Greenleaf manzanita (Arctostaphylos patula)- This ericaceous shrub formed the lower but denser and greater cover of two woody layers in this forest range vegetation. Good natural history of this species was provided by Mozingo (1984, ps. 123-128). Short descriptions but good figures of this species were given in Dayton (1931, p. 124-125), Forest Service (1940, B17), and Sampson and Jespersen (1963, p. 122-124). The latter authors offerred a terse assessment regarding value of manzanita browse: "No manzanita species provide high-quality browse or a large volume of season-long feed". Greatest value of greenleaf manzanita on this range was likely that of watershed protection.

Snake Range, Great Basin National Park, White Pine County, Nevada. June.

34.Quakies in the Canadian zone- In the famous--and still useful--scheme of C. Hart Merriam the Canadian Life Zone was the next highest in elevation and the one above the Transition Zone. Classically, in the mountains of southwestern North America the Canadian Life Zone was characterized by forest communit(ies) consisting of several species of conifers such that it was often regarded as a mixied conifer type. Douglas-fir and white fir tend to dominate with Engelmann spruce and limber pine from forest communities of still higher elevation being associate or subordinate species. Quaking aspen (Populus tremuloides) of the Salicaceae (willow or poplar family) is the only angiosperm tree that achieves dominance among the tall gymnosperm representatives.

Alpine range on the far mountain (Wheeler Peak) was obvious in this Great Basin panorama.

This landscape-scale view of a quaking aspen forest community was at the upper elevtional limits of quakies in the Snake Range of the Great Basin. The forest cover type above (next one up and higher in elevation) the quaking aspen forest was the forest community dominated by Engelmann spruce (Picea engelmannii). Several individual trees of Engelmann spruce were clearly visible in this stand or colony of quakies.

Snake Range, Great Basin National Park, White Pine County, Nevada. June. Elevation of aspen groves ranged from roughly 9,000 to 9,500 feet. Two different communities of quaking aspen were presented in the following photographs so units for ecosystems, Kuchler "vegetation types" forest and rangeland cover types, and other designations were cited in those captions.

35.Grove with a woody carpet- In the Canadian Life Zone of the Snake Range quaking aspen formed a grove with greenleaf manzanita as a shrub or lower woody layer and with much recruitment of young quakies. Most of these are undoubtedly clonal shoots from the creeping woody rootstocks of this colony forming species (ie. most of the "trees" are actually limbs arising from a spreading underground shoot which is the trunk of one genetic individual; the grove is one or a few trees). Quaking aspen recruitment has been primarily by asexual means ("tillering" by trees so to speak) rather than establishment from seed. This method of vegetative regeneration allowed aspen to emerge and co-exist with greenleaf manzanita. Young quakies came up in the middle of dense mats of manzanita so that this range plant community consisted of two species and two layers of vegetation.

This understorey contrsted sharply with that of the more common quaking aspen-dominated herbaceous vegetation that grew adjacent to this aspen-manzanita woody plant community.

Successional status of quaking aspen remains a matter of interpretation. In absence of periodic fire coniferous species (eg. Englemann spruce on the range considered here) successfully invades aspen groves prompting numerous workers to declare the quakies as seral vegetation. Other workers have insisted that fire is a natural part of climate such that when "normal" climate (ie. that which "spawns" fire) is not overridden by fire suppression quaking aspen does fine, thank you. Grazing/browsing of aspen-dominated range vegetation constitutes another natural component of forest and woodland ecosystems in Great Basin, Rocky Mountain, Cascade, etc. ranges that must be included in any analysis of plant succession.

To rangemen, foresters, and wildlifers quaking aspen is a native or natural plant community that is forest and rangeland. This range type provides browse, cover, general habitat, watershed vegetation, aesthetics and recreational value so it merits recognition as both a forest cover type by the Society of American Foresters (Eyre, 1980. ps. 96-97) and a rangeland cover type by the Society for Range Management (Shiflet, 1994, ps. 50-51). Incidentially, both of the authorities just cited stated clearly that quaking aspen was either climax or seral depending on habitat (range or forest site).

Snake Range, Great Basin National Park, White Pine County, Nevada. June, early estival aspect. Elevation was about 9,000 to 9,500 feet. FRES No. 19 (Aspen-Birch Forest and Woodland Ecosystem). There was no meaningful Kuchler unit for western quaking aspen. Many workers, including Kuchler (1964, 1966), interpreted quaking aspen as seral to other "vegetation types" such as Englemann spruce in this case so that quakies did not get a designation/description which, for example, in terminology of Kuchler (1964, 1966) was potential natural vegetation. SAF 217 (Aspen); SRM 411 (Aspen Woodland). Would have to be lumped with Rocky Mountain Montane Conifer Forest 122.6, Douglas Fir-White Fir (Mixed Conifer) Series 122.61, Populus tremuloides subclimax Association 122.614 of Brown et al. (1998, p. 37). Populus tremuloides / Arctostaphylos patula woodland which was not shown either Bougeron et al. (29 August, 1994) or Nevada Natural Heritage Program (26 September, 2003) even when both gave other Populus tremuloides associations "by the droves". Central Basin and Range- High Elevation Carbonate Mountains Ecoregion 13e (Bryce et al., 2003).

36. Invaded but out-distancing the invasion (maybe)- Quaking aspen grove in the Snake Range of Great Basin that was being invaded by Engelmann spruce from the Engelmann spruce forest of the Hudsonian Life Zone just "up the hill". There was recruitment of both Engelmann spruce and quaking aspen with both tree species represented by several age classes. Both species were also represented by dead and downed timber indicating that both had co-existed for several generations. Attention was drawn to the log of Engelmann spruce in right background of second photograph (we will stumble across this log in the next immediately succeeding photograph).

The understorey appeared bare, especially on the predominately east slope presented in the first slide. These photographs were of early summer vegetation from which the last snow had melted only recently and on which there had been grazing by native ruminats. Young plants of the following grasses were present: Cusic bluegrass (Poa cusickii ssp. epilis) , Letterman needlegrass (Sipa lettermanii), fringed brome (Bromus ciliatus= B. richardsonii), and California or mountain brome (B. carinatus= B. marginatus). Blackroot or kobresia sedge (Carex elynoides) and, in smaller amounts, dark or blackened sedge (C. atrata= C. heteroneura) were also present in herbaceous understorey

FRES No. 19 (Aspen-Birch Forest and Woodland Ecosystem). No K- units for what Kuchler (1964, 1966) apparently interpreted as seral and not potential natural vegetation or else could not map at his mapping scale. SAF 217 (Aspen);SRM 411 (Aspen Woodland). For Brown et al. (1998, p. 37) units would have to lump Great Basin with Rocky Mountain (as was done for Subalpine Conifer Forest) which gave Populus tremuloides subclimax Association 122.614 of Douglas Fir- White Fir (Mixed Conifer) Series 122.61 under Rocky Mountain Conifer Forest 122.6. Populus tremuloides / Bromus carinatus Forest (Nevada Natural Heritage Program, 26 September, 2003). Central Basin and Range- High Elevation Carbonate Mountains Ecoregion, 13e (Bryce et al., 2003).

37. Aspen grove with a story but not much understory- Up-to-the-log, close-up view of range vegetation in the quaking aspen woodland shown in the immediately preceding slide. This fallen log (not all logs in such forests and woodlands fall intact) of Engelmann spruce was evidence that this species and quaking aspen had lived together on this forest range site for "many snows". Here in the Canadian Life Zone aspen was replacing itself aminst a corpse of the tree species that is the dominant of the climax Engelmann spruce-limber pine forest of the next vegetation and life zone. Engelmann spruce was also reproducing (sexually in contrast to asexual reproduction of aspen) on this site as was shown in the two preceding photographs.

Grasses and grasslike plants on this range site (more obvious in the second of these two slides) included Cusick's bluegrass, fringed brome, Calirornia or mountain brome, Letterman's needlegrass, blackroot sedge, and dark or blackened sedge. A few miscellaneous forbs were not identified.

Snake Range, Great Basin National Park, White Pine,County, Nevada. June, early estival aspect. Elevatrion was about 9,000 to 9,5000. Ecosystem, forest and rangeland cover types, biotic community, associations, ecoregion, etc. of this range vegetation were cited in the immediately preceding caption.

38. Don't tremble 'til the shutter clicks- Details of quaking aspen presented at the scale of a limb in crown of mature tree (first photograph) and leaves on twig (second photograph). Arrangement and shape of leaves (including curved petiole) of quaking aspen causes these relatively small leaves to twist in the slightest breeze giving the appearance of trembling or quaking. Hence origin of both common and scientific names of this valuable forest and range plant. From a range perspective quakies are valuable as browse plants for just about any animal that can reach leaves, buds, twigs, or even bark.

For general introduction and natural history to quaking aspen Lanner (1984, p. ps. 132-137) was recommended. Old "stand-bys" for treatment of quaking aspen as a range browse plant included Day (1931, p. 17), Forest Service (1940, B111 ), and Sampson and Jespersen (1963, p. 51).

Snake Range, Great Basin National Park, White Pine County, Nevada. June.

39. In the Hudsonian and Engelmann spruce- Engelmann spruce-dominated forest, with limber pine and white fir as minor to associate species locally, in the Snake Range and the lower Hudsonian Life Zone. In this forest Engelmann spruce were substantially smaller than many of the trees in the Rocky Mountains. Soils in the Snake Range are generally rocky and relatively shallow (just wait until we get to the glacial moraine of these hills) and far from optimal for this conifer.

Forest vegetation was primarily a single tree-layer except where fallen trees left gaps that were filled by seedlings and saplings of the Tolerant (Wenger, 1984, p. 3) Engelmann spruce. Understorey--woody or herbaceous--was non-existant to sparse so this was 4-40 (cows need mouths four feet wide and to run 40 miles an hour to survive) range even more than the Great Basin Desert. In fact, here range brutes need to be able to handle high elevation conditions (elevation was 10,000 feet or more). There were some herbaceous range plants that were introduced below. Subalpine fire (Abies lasiocarpa) was absent from this forest vegetation.

Great Basin National Park, White Pine County, Nevada. June. Elevation was 10,000 feet. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-20 (Southwestern Spruce-Fir Forest), basically determined from Kuchler (1964, p. 21) description with dominant Englemann spruce and component species of limber pine, bristlecone pine (Pinus aristata), common juniper (Juniperus communis), and quaking aspen which were in adjoining forest cover types. SAF 206 (Engelmann Spruce-Subalpine Fir). Engelmann Spruce-Alpine Fir Series 121.31 of Rocky Mountain and Great Basin Subalpine Conifer Forest 121.3 of Brown et al. (1998, p. 36). National Vegetation Classification for Nevada (Nevada Natural Heritage Program, 26 Septermber, 2003) did not include a single Picea engelmannii unit (alliance, association, or even a "go-to-hell")! The preliminary National Vegetation Classifiction for Utah and Nevada (Bourgeron et al., 29 August, 1994) gave a Picea engelmannii Alliance, but no associations matched. This was Picea engelmannii Woodland. Central Basin and Range- High Elevation Carbonate Mountains Ecoregion, 13e (Bryce et al., 2003).

40. Up and down-Two samples of range vegetation from an Engelmann spruce climax closed canopy forest in the Snake Range of the Great Basin. Limber pine and white fir were present in this forest. Their cover and density varied from that of associate to incidental species for limber pine and occasional presence of white fir. There was an extremely sparse (usually non-existant) herbaceous understorey, but locally some grass and sedge species formed tiny patches of herbage in a matrix of coniferous crown cover. These species included blackroot sedge, Letterman's needlagrass, California or mountain brome, and fringed brome. There were also spots of pteridophytic alpine or western spikemoss (Selaginella watsonii).

Engelmann spruce has been determined to be a Tolerant species (Wenger, 1984, p.3). Openings formed by death of an existing tree and resultant opening in forest cnopy resulted in recruitment of Engelmann spruce. This phemenon was shown in foreground of the second of these two slides.

The bark of Engelmann spruce in Great Basin forests has a different color (much lighter) and texture from that of Rocky Mountain Engelmann spruce.

Great Basin National Park, White Pine County, Nevada. June. Elevation was 10,000 feet. FRES No. No. 23 (Fir Spruce Forest Ecoosystem). K-20 (Southwestern Spruce-Fir Forest) as was explained in preceding caption. SAF 206 (Engelmann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series 121.31 of Rocky Mountain and Great Basin Subalpine Conifer Forest 121.3 of Brown et al. (1998, p. 36). Closest thing in National Vegetation Classification System (Bourgeron et al., 29 August, 1994) was Picea engelmannii Alliance (without matching associations). Was a Picea engelmannii Woodland. Central Basin and Range- High Elevation Carbonate Mountains Ecoregion, 13e (Bryce et al., 2003).

Forest vegetation at these elevtions in this part of the Great Basin Snake Range occurred as a continuum or gradual transition of forest communities. These different forests ranged from the "pretty much pure" form of Engelmann spruce forests (in which limber pine was an incidental to associate species and white fir was incidental) to a mixed forest of Engelmann spruce-limber pine-white fir forests to limber pine forests to limber pine -bristlecone pine forests. There were also gradual changes in species composition of forest understories with change and most development (such as it was) of understorey in limber pine-bristlecone (timberline) forests.

41. Tree cover and of grass, too- Lower limbs of Engelmann spruce and a nice specimen of California or mountain brome (Bromus carinatus= B. marginatus). This was the most prevalent or abundant (relatively speaking; it was scarce in absolute terms) herbaceous plant in this tract of Engelmann spruce climax forest in the Snake Range of Great Basin. California or mountain brome was also the most abundant herbaceous species in adjoining quaking aspen woodland.

Great Basin National Park, White Pine County, Nevada. Elevation was 10,000 feet. June, this specimen of brome was in pre-bloom stage.

42. Mountain or California brome (Bromus carinatus = B. marginatus)- Mountain brome is one of the most valuable and most productive bromegrass species in Great Basin forests of upper elevations (Canadian and Hudsonian Life Zones), especially quaking aspen woodland, Engelmann spruce- and Engelmann spruce-limber pine transition forests. It was frequently the most frequent or abundant herbaceous species in the almost-absent understorey of these range forest cover types.

Second photograph showed panicle of this plant.

Mountain or California brome is one of the most important native bromegrasses throughouut its range. Forest Service (1940, G33) treated this as a complex of up to three species. Hitchcock and Chase (1951, ps. 31, 35-37) treated B. carinatus and B. marginatus as distincet species but "allies" and noted that these "are relished by all classes of stock" with sheep and horses being quite fond of the panicles. Arnow in A Utah Flora (Welsh et al., 1993, p. 799) treated these two taxa as conspecific and retained the scientific as Bromus carinatus. Arnow explained that B. carinatus is "facultatively cleistogamous" and part of a polymorphic complex of closely related taxa. While mountain or California brome is palatable it is a relatively short-lived perennial or, according to Hitchcock and Chase (1951, p. 35) sometimes a biennial. Stubbendieck et al. (1992, ps. 158-159) incorporated the above conclusions in their description of this valuable native bromegrass that is one of 200 range species on the Society for Range Management intercollegiate range plant contest.

Snake Range, Great Basin National Park, White Pine County, Nevada. Elevation was 10,000 feet. June, this specimen was in early bloom (panicle not fully expanded).

43. Dark, black, or blackeneed sedge (Carex atrata= C. heteroneura)- This grasslike range plant was one of the more common herbaceous species in quaking aspen woodland and Engelmann spruce forests in the lower part of the Hudsonian Life Zone. Blackened sedge formed localized, fairly dense populations along streams in these range cover types. In such mesic, fairly well-lite microsites this sedge was the dominant herbaceous species, but it was absent from drier and shadier habitats.

Mountain brome (presented immediately above) was the overall most common herbaceous plant in Hudsonian Zone forests.

Snake Range, Great Basin National Park, White Pine County, Nevada. Elevation was 10,000 feet. June, anthesis (and extremely attractive).

44. A hard country and hard-looking trees- An example of a limber pine forest cover type that had developed at about the lowest elevational level (roughly 10,200 feet) at which this species becomes dominant in this area of the Snake Range. Engelmann spruce (dominant of the conifer forest that develops at slightly lower elevations), white fir, and common juniper (Juniperus communis) were associate species to limber pine on this "hard-scramble" site. At slightly higher elevations and rockier ground (hard to believe but stay tuned) common juniper becomes associate and forms a lower conifer shrub element. This was evident as the lower, spreading shrubs, especially in right midground.

This erie stand of mixed conifers was the first outpost of limber pine dominance in the subalpine zone (upper level of Hudsonian Life Zone).

Great Basin National Park, White Pine County, Nevada. June, early estival aspect (and still winter was not that far away; winter is never far away at this location and elevation). No FRES unit: Forest Service (Garrison et al., 1977) did not give a forest and range ecosystem designation for limber pine-bristlecone forest. Kuchler (1966, p.22) did give a "vegetation type" for this subalpine potential natural vegetration: Great Basin Pine Forest (Pinus). SAF 219 (Limber Pine): Society of American Foresters (Eyre, 1980) distinguished the limber pine forest cover type from the bristlecone forest cover type. Pinus flexilis / Juniperus communis Woodland (Nevada Natural Heritage, Program, 26 September, 2003). Bristlecone Pine-Limber Pine Series 121.32 of Rocky Mountain and Great Basin Subalpine Conifer Forest 121.3. Central Basin and Range- High Elevation Carbonate Mountains Ecoregion, 13e (Bryce et al., 2003).

45. Limber pine forest with a taste of white fir- On a slightly more favorable local habitat white fir served as an associate species in a limber pine forest found just above the Engelmann spruce-dominated forest zone. The foremost sampling in right foreground was a white fir that could be readily identified from the "blisters" of pitch or rosin on its trunk.Seedlings in extreme left foreground were limber pine.

Great Basin National Park, White Pine County, Nevada. Elevation was about 10,200 feet. June, early estival aspect. Designations of Kuchler "vegetation type", forest cover type, biotic community, association, ecoregion, etc.were cited in the immediately preceding caption.

46. Old limber pine with common juniper at its feet- A mature limber pine with characteristic bark features consistent with its age was growing on a more favorable microhabitat in the limber pine-Engelmann spruce transition forest. A common juniper was growing in the shade of the pine. Note limber pines in background with most of crowns missing, blown-off perhaps.

Snake Range, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,200 feet. June, early estival aspect. Designations of forest cover type, Kuchler "vegetation type", biotic community, association, ecoregion, etc. were cited in the immediately succeeding caption.

47. Dominant and associate of the limber pine range type- Forest Service (Garrison et al., 1977) did not list the limber pine type in any forest or range ecosystem. Kuchler (1966, p. 22) "vegetation type" was Great Basin Pine Forest (Pinus). SAF 219 (Limber Pine). Bristlecone Pine-Limber Pine Series 121.32 of Rocky Mountain and Great Basin Subalpine Conifer Forest 121.3 of Brown et al. (1998, p. 36). Pinus flexilis / Juniperus communis Woodland (Nevada Natural Heritage Program, 26 September 2003). Central Basin and Range- High Elevation Carbonate Mountains Ecoregion, 13e (Bryce et al., 2003).

48. Scrub forest on talus of quartzite- Landscape-scale view of part of the Snake Range with limber pine forest in foreground that developed on glacial moraine composed primarily of quartzite. Quartzite is an extremely hard metamorphic rock derived from an original sandstone that was high in quartz (ie. quartzite is metamorposed quartz sandstone; a metamorphosed sedimdntary rock). Most quartzite is over 90% quartz. According to Wikipedia encyclopedia (Wikimedia Foundation) quartz sand grains and quartz silica cement fused when the original sandstone was heated and pressed and brought into contact with magma by tectonic forces. This resulted in a rock material of interlocking quartz grains so that quartzite breaks through rather than among quartz grains. Thus quartzite does not disintegrate into small stones or grains of quartz but is typically found as stones with smooth rather than sharp edges even though these rock have are angular with numerous facets (faces).

Quartzite at this location near base of Mount Wheeler was on a glacial moraine which is a "distinct accumulation of unsorted, unstratified glacial drift, predominately till, deposited chiefly by direct action of glacier ice ..." (Gary et al., 1972). The deep, piled deposit of quartzite along this mountain side had later formed or was arranged on (at least closely resenbling or functioning as) a talus slope. Talus was defined by Gary et al., 1972) as: Rrock fragments of any size or shape (usually coarse and angular) derived from and lying at the base of a cliff or very steep, rocky slope. Also, the outward sloping and accumulated heap or mass of such loose broken rock, considered as a unit, and formed chiefly by gravitational falling, rolling , or sliding". Talus slope refers to "a steep, concave slope formed by an accumulation of loose rock fragemens; especially such a slope at the base of a cliff or steep slope and formed by the coalescence of talus cones; the surface profile of an accumulation of talus" (Gary et al., 1972).

This quartzite forest site is undoubtedly one of the most unfavorable environments for vascular rangeland and forest plants. Unlikely as it would seem, various shrubs and herbaceous plants grew on this harsh habitat along with limber pine. Photographs presented below showed some of these plant species. This slide gave a general view of this range plant community.

There were a few Great Basin or western sbristlecone pine (Pinus longaeva) in this limber pine forest, especially in extreme right background. The bristlecone pine-limber pine forest (more like woodland) community was featured in a series of slides below.

No FRES unit: Forest Service (Garrison et al., 1977) did not give a forest and range ecosystem designation for limber pine-bristlecone forest. Kuchler (1966, p.22) did give a "vegetation type" for this subalpine potential natural vegetration: Great Basin Pine Forest (Pinus). SAF 219 (Limber Pine): Society of American Foresters (Eyre, 1980) distinguished the limber pine forest cover type from the bristlecone forest cover type. Pinus flexilis / Juniperus communis Woodland (Nevada Natural Heritage, Program, 26 September, 2003). Bristlecone Pine-Limber Pine Series 121.32 of Rocky Mountain and Great Basin Subalpine Conifer Forest 121.3. Central Basin and Range- High Elevation Carbonate Mountains Ecoregion, 13e (Bryce et al., 2003).

49.The next generation coming through the talus- A seedling of limber pine had germinated and was growing on a talus slope of quartzite at base of Mount Wheeler in the Snake Range. Great Basin National Park, White Pine County, Nevada. Elevation was about 10,200 feet. June.

50.Simplier plants of harder country- Alpine or western spikemoss (Selaginella watsonii) thriving on quartzite glacial moraine in the Snake Range of the Great Basin. The spikemosses are some of the most primitive vascular plants in the Great Basin or surrounding provinces. Spikemosses are those plants known as pteridophytes or vascular cryptogams. Cryptogams are those plants which reproduce by spores or gametes rather than seeds and include algae, bryophytes, and pteridophytes. The latter (division of Pteriophyta) are thus spore-bearing vascular plants like spikemosses, horsetails, and ferns.

Cryptogams are important in soil-forming processes and are some of the first plants to appear on a sere.

Great Basin National Park, White Pine County, Nevada. Elevation was about 10, 200 feet. June.

51. Letterman's needlaegrass (Stipa lettermanii)- This grass species produced some of the largest herbaceous plants in the understorey of limber pine forest vegetation and, especially, in limber-pine-Engelmann spruce transition forests. This is not a major Stipa species on the Western Range, but on forest ranges with understories as scantily covered as contemporary college co-eds this species is extremely valuable and pleasing to behold, though perhaps not as much so as other sacnty items. (Rangemen have to take their pleasures where they find them.)

Note cone of limber pine in front of needlegrass carefully included for readers' education and viewing pleasure.

Snake Range, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,200 feet. June.

52.Details of Letterman's needlegrass- Shoots of Stip lettermanii complete with spike inflorescences.

Snake Range, Great Basin National Park, White Pine County, Nevada. Elevation was about 10, 200 feet. June.

53. Fringed brome (Bromus ciliatus= B. richardsonii)- This species is one of the most widely distributed native perennial bromegrasses in North America. Its range extends from Newfoundland to British Columbia and Alaska and south to Texas and Mexico.

Forest Service (1940, G34) wrote that this bunchgrass "ranks with the choicest forage grasses in palatability" and that it was "... relished by all classes of livestock". Arnow (in Welsh et al., 1993, 800) noted: "fringed brome is highly palatable to all classes of livestock and to deer and elk. Although it does not form dense stands, the species is sufficiently common to constitute a moderately good aource of forage on some forest ranges".

The "nodding" panicle is characteristic of this but also several related Bromus species. This panicle was just beginning to spread its secondary branches off of the rachis.

Snake Range, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,200 feet. June, early bloom stage.

Farther up in the Hudsonian Life Zone there is the final (highest elevation) belt of forest or woodland known as the subalpine, the uppermost limit of which is timberline or treeline. (Alpine ecosystem is the upper elevational limit of vascular plant life and range vegetation.) In the Snake Range the upper outpost of woody range vegetation developed at slightly higher elevations than (or the upper end of ) limber pine forests. It was at this elevation that western or Great Basin bristlecone (Pinus longaeva) began to appear. At this location near Wheeler Peak western bristlecone pine does not form "pure" stands but rather grows as co-dominant with limber pine. Both of these species grow and survive to great ages, especially bristlecone pine, on the quartzite glacial moraine forest site. Limber pine is classified as Intolerant and bristlecone pine is Very Intolerant (Wenger, 1984, p. 3). These two Pinus species cannot compete with other trees and basically have the most inhospitable, harshest sites that "nobody else wants" all to themselves (at least tree-wise).

It is on the barren, wind-swept, arid pile of quartzite rocks that bristlecone grows to their almost immortal ages. Someone astutely observed that "it is not that bristlecone pine lives so long as much as it that it takes so long to die". Bristlecones on hostile habitats grow extremely slow (perhaps adding an inch of trunk diameter in a century) and their needles persist for numerous growing seasons. These specimens are prized for their dendrochronological, especially dendroclimatological, records. "Thereon hangs a tale" (see Cohen, 1998).

The silvics of bristlecone pine was not covered in Burns and Honkala (1990), but they did treat limber pine (Burns and Honkala, 1990, ps. 348-354). A readable natural history of the western bristlecone pine is that of Lanner (1983, 24-29). A longer, more detailed though still not highly technical account of the bristlecone pine is that of Cohen (1998). One of the older, popular, and cultural accounts of bristlecone pine is that of Peattie (1959, 58-62) who, appropraiately enough, dealt first with limber pine (Peattie, 1959, ps. 34-37).

The western or Great Basin bristlecone pine (P. longaeva) was distinguished from the Rocky Mountain bristlecone (P. aristata) by dendrologists. For good, readily available discussions see Harlow et al. (1979, ps. 76-77) and Welsh et al.(1993, p.35).

It is useful to follow Harlow et al. (1979, ps. 61, 71-77) and point out that limber pine is designated as a stone pine whereas bristlecone pine is a foxtail pine (and pinyon pines are nut pines) with all being in the soft pine group (subgenus: Haploxylon). Stone and nut pine refers to the large, edible seed whereas foxtail obviously draws attention to the full, bushy appearing needles of bristlecone pine branches. Both limber and bristlecone pine have five needles per fascicle.

Short, but easily gotten summary of the subalpine limber pine and bristlecone pine woodland is that of Vasek and Thorne (in Barbour and Major, 1995, ps. 823-826).

The series of photographs of a Great Basin bristlecone pine-limber pine forest presented below were of the famed Wheeler Peak grove in Great Basin National Park (formerly Humboldt-Toiyabe National Forest). Emphasis was on the natural vegetation--as range of course--and not "gee-whiz", "look at them neat ole trees" photographs from color-enhancing film and filter.

54. Yes, that is a forest. (Would you believe a woodland?)- Great Basin bristlecone forest with limber pine as co-dominant and common juniper as associate species. This subalpine woodland developed on the quartzite glacial moraine forest site, one of the harshest of all habitats for trees. Characteristic gnarled, twisted, rust-colored trunk and limbs of an ancient western bristlecone was obvious and the stuff of photographic legend.

Common juniper constituted a lower woody (shrub) shrub layer. The other shrub in this pine woodland was gooseberry currant (Ribes montigenum= R. lacustre var. lentum).

There was a very sparse understorey which was treated at end of this series below.

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10, 400 feet. June. No FRES unit: Forest Service (Garrison et al., 1977) did not give a forest and range ecosystem designation for limber pine-bristlecone forest. Kuchler (1966, p.22) did give a "vegetation type" for this subalpine potential natural vegetration: Great Basin Pine Forest (Pinus). SAF 209 (Bristlecone Pine): Society of American Foresters (Eyre, 1980) distinguished the limber pine forest cover type from the bristlecone forest cover type.Bristlecone was chosen here based on elevation and diagnostic presence (indicator species) of Pinus longaeva in this forest or woodland community. Pinus longaeva - Pinus flexilis Woodland (Nevada Natural Heritage, Program, 26 September, 2003). Bristlecone Pine-Limber Pine Series 121.32 of Rocky Mountain and Great Basin Subalpine Conifer Forest 121.3. Central Basin and Range- High Elevation Carbonate Mountains Ecoregion, 13e (Bryce et al., 2003).

55. Pines on quartzite- Western bristlecone and limber pines typically grew in close proximity (often "cheek-by-jowl" or would it be "snag-by-bole"). In the first of these two slides two bristlecone pines flanked on both left and right an old, broken limber pine with a young trunk while a third bristlecone pine was growing behind (and slightly to the right) of the limber pine. The latter had somehow lost most of its trunk and crown, but upon being released from apical dominance a lower shoot had become the new trunk of the old tree.

In the second of these two slides a still-living but mostly bare snag of an ancient bristlecone pine (center-most tree; bark-peeled trunk) had two smaller, younger bristlecone pines immediately to its left. The next (and larger) tree to the left was a limber pine. Three smaller trees (three distinct trunks) to the far right were limber pines. The pattern of size and number (density) of limber pine and bristlecone pine shown in these two slides was typical of this bristlecone-limber pine woodland vegetation. Limber pine typically outnumbered bristlecone pine, but the latter had larger (probably older) trees and thus greater crown cover. On criteria or inherent definition of forest cover (= dominance) types bristlecone pine "got to name the type" for this woodland range vegetation.

Students should observe that it was very easy to distinguish between limber and bristlecone pines. The "can't miss it" foxtail appearance of bristlecone, the foxtail pine, was unmistakeable and contrasted readily with the more typical pine boughs of limber pine. For that we move to the next slide...

56. Tree full of foxtails - Textbook shot for students new to bristlecone pine. Characteristic bark of a mature bristlecone plus the namesake foxtail-appearing arrangement of dense needles on bristlecone branches. In side-by-side comparisons bristlecone and limber pine cannot be confused. A blind forester could tell the difference-- as long as he could feel of the branches and needles or of the bark.

Shrubs in understorey ( left margin; center midground between two tree trunks) was common or spreading juniper.

57. In it together up to their crowns- An ancient bristlecone pine (left and slightly behind) and an immense limber pine (right and to somewhat to the forefront) were dancing bark-to-bark on a quartzite moraine in upper limits of the Hudsonian Life Zone. Both pine species had basal boughs that formed a local lower woody layer. These lower branches and seedlings of the two species sometimes joined common or spreading juniper and gooseberrry currant as part of a second or lower woody strata of this conifer woodland range. This was the upper elevtional limit of subalpine woody vegetation.

It was left to students to decide who was leading whom (ie. which species was leading which). Perhaps that was a question in regards to plant succession and seral species.

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10, 400 feet. June.

58. Interior of a bristlecone-limber pine wodland range- Co-dominant limber pine (adult tree immediately left of plaque) and bristlecone pine (adult tree immediately behind and slightly to right of plaque). The National Park Service thoughtfully plaaced this educational plaque in the center of Wheeler Grove subalpine pine woodland to help visitors lear to identify these two co-dominant Pinus species. Bristlecone pine is one of the so-called foxtail pines. Bristlecone branches have needles along their internodes which then have the "bushy tail" appearance. By contrast limber pine has the more common characteristic of bare internodes with needles limited to more terminal parts of branches. Both species have five-needle fascicles (ie. leaves are in units of five).

The other key field-identifiction feature useful in distinguishing these Pinus species in the bark. Mature bark of limber pine is moderately to deeply furrowed and with a pattern of relataively large, oblong plates. Mature bark of bristlecone is more smooth with smaller and more square bark plates.

Possible point of confusion: many of the very old trees ("ancient" was not an overworked adjective in describing trees of this grove) of both pine species have lost much of their bark thereby exposing the smooth wood beneath. This wood has been even more "sanded" by windblown particles of snow, ice, soil, small rock fragments, etc. over centures of exposure. Such smoothed wood in combination with the odd-shaped trunks and limbs of old trees results in the driftwood-like appearance which is the "tradmark" of the bristlecone pine (of course many such trees are limber pines). Newcomers should not confuse weather-worn wood with bark.

Limber and bristlecone pine often have persistent lower limbs coming directly off of the lower trunk almost at ground level. This feature (which also was shown in the next two preceding slides) results in the two dominant trees forming a continuous woody layer from near soil surfact to crown. Wood and leaves of bristlecone and limber pine along with common juniper and gooseberry currant comprise a lower woody layer of thei woodland range vegetation. Common juniper and current were present in this understorey but were not descernible in this slide. Clearly visible, however, were young tips of branches with conspicuous light green clusters of new needles. Grass and sedge species were also growing in this pine grove so as to give an herbaceous component (not exactly a layer) to this woodland plant community. These other species of the subalpine pine woodland were presented below.

59. Bristlely patriarchs on hard rock- Local (small-scale) stand of bristlecone pine growing on the quartzite, talus slope-resembling glacial moraine of the Snake Range (Wheeler Peak was partly visible in the background). All trees were bristlecone pine except for young tree on far right and tree in center background). Very little understorey except for gooseberry currant and some grasses and grasslike plants (none of this were visible in this photograph) plus lower living limbs which frequently arise from basal portions of trunks even in very old trees of both Pinus species. This feature of having live low limbs almost at ground level was visible in the photograph immediately preceding and the photograph immediatedly succeeding this slide.

The large bristlecone pine in left foreground was an example of the "trademark" driftwood-like appearance of very old trees of both pine species, especially bristlecone pine.

60. Youngsters amid the rubble of quartsite and ancestorial remains- Growing on talus slope-resembling glacial moraine in the minst of quartzite rock and dead pine corpses (both standing and down) were relatively young bristlecone pines. In the first of two slides a bristlecone pine stands remarkably tall and straight against a partial backdrop of Wheeler Peak. This straight bole of this tree gave an example of "middle-age" bristlecone pine bark. It also showed the common characteristic of survival of the first (oldest) limbs of bristlecone well into maturity. It was described above how these lower limbs that come directly off of the basal trunk make up part of the lower woody layer (along with common juniper and gooseberry currant) of the limber pine-bristlecone pine woodland. The three trees in foreground of this first photograph were bristlecone pine and each was an example of persistence of lower limbs as well an the presence of needles along internodes of branches creating the "bushy tail " appearance of this specis of foxtail pine.

In the second of these slides young bristlecone pine showed the bush-like habit characteristic of this age tree. The lower branches often persist into adulthood and remain part of a lower woody strata of this woodland range vegetation. In extreme right margin of this slide the branch of a limber pine was visible which can be contrasted with those of the foxtail bristlecone pine. Also in in this second photograph was a well-developed local patch of herbaceous vegetation that survived under protection of the lower limbs of bristlecone pine.

Regeneration of the same species of plant in the shade of its parents is often taken as a characteristic of climax species (ie. they replace themselves instead of preparing the sere for other and succeeding plant species). Bristlecone pine is a Very Intolerant species (Wenger, 1984, p. 3), but this quartzite forest site is so harsh that other treee species cannot survive (=invade). Limber and bristlecone pine continue to invade or reproduce their species (recruitment of pine continues indefinitely in successional time scale).

61. Standing guard at the top- Co-dominant bristlecone pine (old, gnarled tree in center foreground plus two young trees at to left of old tree with characteristic foxtail branches) and limber pine (tallest tree and behind gnarled trunk). Silhouetted branches of the two species contrasted features of pine leaders (branches).

It should be observed that young bristlecones had germinated and grown so close to pines of the older generation as to almost be in contact. Generational replacement of the same species is one characteristic of climax plant species.Shelford (1963, p. 163) stated that the bristlecone pine and limber pine vegetation of the Rocky Mountains wa an "edaphic climax on dry rocky slopes and windswept ridges". The current author concurred and reached the same conclusion for these species in Great Basin mountain ranges. Limber pine and bristlecone forest cover types are the climax plant community, the potential natural vegetation on which to base range management and related natural resource decisions.

62. How much and how many and still it grows- Along this footpath through the Wheeler Peak grove of bristlecone pines an ancient and large specimen gave testament to the remarkable longevity and unique anatomy and physiology of it species. Much of the bark of this tree had been lost and it would be supposed that such exposed wood was dead. At other locations including base of the trunk new growth appeared (or very old growth persisted). Apparently intercalary meristem is capable of growth at remarkably old age (or new meristem is formed or regenerated into old age).

What stories such trees could tell, but then how much in the way of "news" ever happened here? This self-guided tour through the "old ones" was an example of the educational and recreational (and in the original meaning of the word) value of natural resources that contribute no commodities or raw material to attend such human needs as food, fiber, shelter, or water. But as John Milton concluded in his poem, Oh His Blindness, "They also serve who only stand and wait". Carry on old tree.

This view was deep in the interior of a limber pine-bristlecone pine grove where bristlecone pine was locally; dominant.

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June.

63. A "coming-out party" for Great Basin gals- Immature female cones of bristlecone pine. This coloration is distinctive and userful for purposes of identifiction.

Wheeler Grove, Great Basin National Prk, White Pine County, Nevada. Elevtion about 10,400 feet. June.

64. A study of bristlecone pine- This photograph of a leader (branch) of bristlecone pine showed: 1) mature female cone with the single bristle at apex of a cone scale, 2) arrangement of five needles per fascicle, 3) presence of needles along intenode responsible for toxtail appearance and 4) apical bud (behind cone).

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10, 400 feet. June.

65. Gooseberry currant (Ribes montigenum= R. lacustre var lentum)- This species was the second most common or major shrub in understorey of subalpine limber pine- bristlecone pine woodland. The most abundant or frequent shrub was common juniper. This gooseberry currrent was growing at base of a very old bristlecone pine. This Ribes species was well-armed with spines. Traditionally Ribes species with spines are known as gooseberries while the unarmed or spineless species are called currants. In the Intrmountain West there are only two or three Ribes species with spines so the term currant is used generically. This species was designated by Welsh et al. (1993, p. 642) with a "hybrid" common name. (Maybe that will please everybody.)

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June.

66. Range on the moraine- Of course there was range of the talus slope-like glacial moraine of quartzite and in the subalpine limber pine-bristlecone pine woodland. It just takes a special critter to range on it. Herbaceous plants were present on the quartzite parent material of the limber pine forest and the limber pine-bristlecone pine forest (woodland was more precise) that was of slightly higher elevation than the "straight" limber pine forest. Several herbaceous species grew well (relatively speaking) on local habitats of shallow soil among the quartzite rock and pines. These species included alpine or Watson's spikemoss, blackroot sedge, spike trisetum (Trisetum spicatum), Cusick's bluegrass, Letterman's needlegrass, and fringed brome .

The little stand of herbaceous range vegetation shown here was deep in the central part of a grove of bristlecone pine. The predominant species here was blackroot sedge. There was also some Cusick's bluegrass.

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June.

67. Blackroot sedge in subalpine woodland- In the upper part of the Hudsonian Life Zone and on quartzite of a glacial moraine a subalpine limber pine-bristleleaf pine woodland had developed. This range type had an extremely limited but distinctive herbaceous component. Blackroot sedge was one of the more common herbaceous species and appeared to be the major grasslike range plant.Note the naturally fallen foxtail twig tip in the second photograph and graphically placed cone in the first photograph, both of bristlecone pine.

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June.

68. Blackroot sedge (Carex elynoides)- Carex is a remarkable genus whose species occupy a wide array of habitats when viewed from standpoints of numerous soil features, elevation, vegetation, and especially water. Some carices are aquatic species whereas others are mesophyes which this latter ranging from those whose habitats are characterized by deep soils of extremely mesic conditions to others that thrive on environments of shallow, rocky, and generally droughtly soils. Blackroot sedge obviously came closer to the latter range environment.

Blackroot sedge is a non-rhizomatous or cespitose sedge. Interestingly this was consistent with dominance of bunchgrasses in Snake Range forest range types. Perhaps habitats in these forest communities that are favorable for grasses and sedges occur at micro-scale (= microhabitats) such that tufted species have a competitive (natural selection) advantage over sod-forming (stoloniferous and/or rhizomatous) species. Blackened or dark sedge of the Engelmann spruce and quaking aspen forest (covered above) is also a tufted perennial without rhizomes. By way of comparison, marshes in the adjoining Snake Valley had as the major Carex black-creeper or silver sedge (C. praegracilis), a species characterized by large, dark, creeping rhizomes (See Hurd et al., 1998, ps. 108-109, 130-131, 208-209).

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June.

69. Local population of spike trisetum on quartzite- This member of the oat tribe appeared to be the major herbaceous species of the limber pine-bristlecone pine aubalpine woodland. No plant species--including pines--were common of the wind-blasted, rock-strewn, shallow soil of this glacial moraine. This range habitat was more like that of a talus slope than typical glacial till. Plants of all species were widely scattered except where 1) young pines grew next to their parents or trees of their parents' cohort and 2) shrubs and/or herbaceous species grew near the dominant pines that provided more favorable microhabitats (eg. spike trisetum and blackroot sedge grew under pine boughs that came off of basal parts of tree trunks).

Observant students would have noticed that the herbaceous plant along left margin of photograph (next to a large rock) was alpine, western, or Watson's spikemoss, a pteridophyte or vascular cryptogam. Spikemoss was not common, but this range plant did occur in understorey of Engelmann spruce forests as well as spruce-limber pine transition forests, limber pine woodlands, and limber pine- bristlecone pine woodlands.(Spikemoss was introduced earlier in this series devoted to elevationally zonal forests/woodlands).

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June.

70. Spike trisetum (Trisetum spicatum)- Two examples of the most prevalent grass in limber pine-bristlecone pine woodland in the Snake Range of Great Basin. This species is one of the 200 range plants on the Society for Range Management intercollegiate contest, an "esteemed position" that bespeaks its importance on the Western Range.

In the first slide alpine or Watson's spikemoss was once again growing beside spike trisetum. This "odd couple" must have had something going because they were more conmonly than not associated species on limber pine-bristlecone pine woodland. These two range plants were growing close to but not directly under a bristlecone pine. The bristlecone cone that had been placed near the plants by the author for instructional purposes had rolled within about 10 to 15 feet of them (and don't call me a *+&#~*^# "nature faker").

Diversity of color in quartzite was obvious in this series of slides.

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June.

71. Cusick's bluegrass or skyline bluegrass (Poa cusickii= P. epilis= P. cusickii ssp.epilis=P. fendleriana) and a Watson's penstemon (Penstemon watsonii= P. phologfilius) for good measure- Several of the Poa species resemble each other enough and/or are variable enough within what was interpreted as a species that taxonomic treatments have varied drastically from one manual or flora to the nest. The speciment shown here was skyline bluegrass that growing on quartzite in a limber pine-dominated woodland at an elevation at which bristlecone pine was just making its appearance. P. epilis= P. cusickii ssp.epilis is usually present as a dioecious species. This specimen was a male. A good field identificatiion feature of skyline or Cusick's bluegrass is the purple-tinged spikelets.

The range forb was Watson's penstemon (as could best be determined from its unopened, "almost-ready" flowers).

Both species of subalpine range plant was in both photographs.

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June, full-bloom for bluegrass and early bloom for penstemon)..

72. Shoots with pancicles in Cusick's or skyline bluegrass- This wind-blown, spreading inflorescence of Cusick's bluegrass had the typical purple-toned spikelets that are a farily reliable characteristic by which to quickly identify this variable species on the range. In the forests and woodlands in the Hudsonian Life Zone of the Snake Range the understorey was very sparse with herbaceous plants being so "few and far between" that such range was by necessity restricted to wildlife. This was especially the case on the glacial moraine covered by quartzite rocks. This quartzite range site remains useful habitat for species like the desert bighorn sheep (Ovis canadensis nelsoni), a native game ruminant the numbers of which declined drastically and are only slowly increasing.

The desert bighorn sheep is the "State Animal of Nevada". Yes, the Silver State does have a State Bird and a State Reptile so one would think that the Nevada legislature would have been astute enough to correctly designate the native bighorn sheep as the State Mammal, but it was politicians who passed this legislation. It would appear that the desert bighorn, whose most critical range is that in mountains like the Snake Range, needs all the help it can get if it is to survive in the Great Basin.

Maybe Cusick's bluegrass (and other range plants) should be left as range for the desert bighorn sheep and associated mule deer. Come to think of it, the bluegrass probably needs all the help it can get too.

Wheeler Peak Grove, Great Basin National Park, White Pine County, Nevada. Elevation was about 10,400 feet. June.

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