Forests, woodlands, and savannas dominated by ponderosa or western yellow pine (Pinus ponderosa) constitute one of the most widespread and, acreage-wise, largest of the forest range types in western North America. The readily read natural history of ponderosa pine by Peattie (1953, ps. 79-86) and the silvics of this species in Burns and Honkala (1990) are always good (and nice) places to start with any tree species, and this is particularily so with ponderosa pine. Rocky Mountain ponderosa pine (P. ponderosa var. scopulorum) is the more easternly of two species of Pinus ponderosa. Rocky Mountain ponderosa pine has a biological range extending from southern Canada to northern Mexico and from northwestern Montana to Trans Pecos Tex through the Black Hills of South Dakota, Nebraska Sandhills and through the mixed prairie in northwestern Okahoma.

Throughout the vast species range Rocky Mountain ponderosa pine occurs at elevations typical for those of Great Plains tallgrass and true prairie range types to lower montane through upper montane forests in mountainous terrain. In this latter elevational zone, western yellow pine most commonly occurs at beginning of forest line above pinyon pine (Pinus edulis)-juniper (Juniperus spp.) woodland or that of the lower montane Douglas-fir (Pseudotsuga menziesii) forest cover type. There are sometimes ecotones among these various range types or, in some unique habitats, transition forest communities where dominant tree species of various forest cover types form a distinctive forest range type (eg. the Rocky Mountain mixed conifer type).

This chapter of Rocky Mountain (and outlying) forests began with treatment of Rocky Mountain forest types dominated and defined primarily (at least in part) by ponderosa pine. There is no other forest tree species that forms such valuable and varied forest range as does ponderosa pine. (This includes both varieties of this species.) In large part this is due to the more open canopy of ponderosa pine range which is a function of the relatively low tree density (= wide spacing of trees) which permits adequate light for growth of herbaceous and shrub species and, therefore, development of lower layers of vegetation. These lower strata in the forest community provide forage and/or browse for range animals.

6. White or pale death camas (Zigadenus elegans)- Pale death camas growing on a shaded hillside in the ponderosa pine forest of the Black Hills. This member of the lily family (Liliaceae) is one of about 11 Zigadenus species native to North America according to recent treatments (Burrows and Tyrl, 2001, ps. 787-792). All of these species should be regarded as toxic, but relatively few pose serious livestock poisoning hazards (Burrows and Tyrl, 2001, p. 790). Death camas is, however, one of the classic stock poisoning range plants and was included here for that purpose. White death camas has a large inflorescence that is quite conspicuous on the floor of the Black Hills ponderosa pine forest.

Students are referred to the classic text-reference by Kingsbury (1964, ps. 461-466) as well as the more recent and encyclopedic Burrows and Tyrl (2001).

Custer State Park, Custer County, South Dakota. July.

7. Forest and grassland- A mosaic of ponderosa pine (Pinus ponderosa) pine forest with an herbaceous understorey dominated by slender wheatgrass (Agropyron trachyculum= A. subsecundum= A. caninum) and true prairie grassland of midgrasses the major species were little bluestem, western wheatgrass, needle-and-thread, and bluebunch wheatgrass (in that overall relative order). The conspicuous forb in foreground of both of these slides was large Indian breadroot, breadroot scurfpea, or prairie-turnip (Psoralea esculenta).

This range vegetation could be seen as either forest with grassland communities developing as glades within or, alternatively, as grassland with covers or groves of forest or woodland. A third perspective or mental/verbal imaging of this Northern Great Plains vegetation was that of a savanna in which groves of ponderosa pine forest comprised a woody component of the overall (at regional-scale) predominant grassland resulting in a savanna physiogonomy. Savannahs have traditionally been regarded as ecotones or transition zones between grassland and woody (tree and/or shrub) vegetation (Dyksterhuis, 1957) This composite or mosaic vegetation did not handily the accepted or classical savanna concept of Dyksterhuis(1957) because these ponderosa pine forest had developed as "vegetational islands" (isolated remnants or, perhaps, relicts of forests that formed over earlier climatic periods) and not as extensions of present era-forests.

Certainly the open canopy, herbaceous understorey, ponderosa pine forests were in monoclimax theory postclimax vegetation (a more mesic habitat-requiring plant community) within the zonal or regional present climax (climatic climax). Polyclimax or climax pattern theories could just as handily provide explanation (successional intrpretations) and description of this patchwork of range vegetation. Obviously groves of ponderosa pine forest developed on different range sites than those of midgrass grassland.

Whatever the visualizatioon or interpretation it was interesting, eductional , inspiring range vegetation. Perhaps even more to the point was the productivity of these various range plant communities and their production of landscape "goods and srvices". For example, a large flock (rafter or gang) of Merriam's wild turkey (Meleagris gallopavo merriami) were foraging at the edge of forest and grassland presented in these and subsequent slides. Unfortunately, the turkeys did not welcome intrusion by this photographer and they vamoosed before the Nikon could capture them in their habitat. Perhaps this was appropriate from the standpoint of nativeness given that Range Types of North America was devoted to native flora. It has been agreed among wildlifers that turkey was not part of the native or indigenous fauna to that part of North American that is now the state of Montana wherein this avian species was introduced by the whiteman strarting in the 1950s (Dickson, 1992, p. 374).

This mosaic of forest and grassland provided a good example of where Landscape Ecology is a more apporpriate framework then Ecosystem Ecology from which to view and analysis range (in this case both rangeland and grazable forest).

Powder River County, Montana. Mid-June; late vernal aspect. Mosaic of FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Grama-Needlegrass-Wheatgrass), SRM 606 (Wheatgrass-Bluestem-Needlegrass), Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

8. At the edge- Margins of ponderosa pine forest with understorey dominated by slender wheatgrass (background) and true prairie grassland of little bluestem, western wheatgrass, needle-and-thread, and bluebunch wheatgrass. The conspicuous and abundant forb in foreground was large Indian breadroot, breadroot scurfpea, or prairie-turnip. Microrelief resulted in a low-lying depression dominated by western wheatgrass (immediate foreground). Adjacent to and above this swale-like low area (immediate right foreground and midground) the grassland was dominated by little bluestem (buff-colored clumps) with needle-and-thread and bluebunch wheatgrass as associates. Needle-and-thread was the most widespread grass with abundant populations in both the western wheatgrass-dominated and little bluestem-dominated grassland communities.

Young ponderosa pine had invaded the grassland (covered in immediately succeeding slide-caption set).

Powder River County, Montana. Mid-June; late vernal aspect. Mosaic of FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Grama-Needlegrass-Wheatgrass), SRM 606 (Wheatgrass-Bluestem-Needlegrass), Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

9. Three zones and woody invasion- A patchwork of climax range vegetation in the unglaciated part of Northern Great Plains. In restricted areas of the northern plains a combination of true prairie grassland and groves or small stands of ponderosa pine-bunchgrass forest had developed. An example of this grassland-forest mosaic was presented in these two slides. The grassland consisted of two major communities: 1) western wheatgrass consociation with needle-and-thread as the associate speciess on low-lying local areas (depressions) and 2) little bluestem-dominated higher areas with needle-and-thread and bluebunch wheatgrass as associate species. Needle-and-thread was the most widespread grass species. In restricted local areas needle-and-thread was co-dominant with little bluestem. An example of the wesstern wheatgrass consociation was in the foreground of both slides. The little bluestem-dominated grassland was in the midground and easily recognized by the light tan- or buff-colored grass clumps. Small saplings and a few seedlings of ponderosa pine were growing on the little bluestem grassland (two paragraphs below in this same caption). Also in both grassland communities were numerous short plants of turnip-root, breadroot scurfpea, or large Indian breadroot.

The ponderosa pine-bunchgrass forest (background of both slides) had developed on the highest ground of this landscape. The forest understorey was exclusively herbaceous with slender wheatgrass as the dominant species. There were some scattered plants of the suffructicose fringed sage or fringed sagebrush in the herbaceous forest layer. The most common forb in the understorey was wavyleaf thistle.

The grassland had been (and was continuing to be) invaded by young ponderosa pines, but this was limited to the little bluestem-dominated community on higher, better-drained ground (observe carefully in both slides). Ponderosa pine typically responds best on well-drained soils (on some forest sites ponderosa pine appears tr require well-aerated, rapidly drained soils). Role of fire in maintaining this grassland and preventing development of ponderosa pine forest in this range landscape was not known to this author. It has likely received only incidental study on this series of range sites. Ponderosa pine is certainly tolerant of surface fire, but even rather low-intensity fire does thin younger pines from forests (even cool fires frequently kill a lot of smaller ponderosa pines) so as to maintain the herbaceous understorey that characterizes ponderosa pine forest across most of the biological range of this species.

Powder River County, Montana. Mid-June; late vernal aspect. Mosaic of FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Grama-Needlegrass-Wheatgrass), SRM 606 (Wheatgrass-Bluestem-Needlegrass), Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

10. Border of two range types- Outermost margin of a ponderosa pine-slender wheatgrass habitat type where it was contiguous with true prairie dominated by little bluestem and needle-and-thread and with bluebunch wheatgrass and western wheatgrass as associate species. Main forb in the grassland was prairie-turnip, large Indian breadroot or breadroot scurfpea. Details of the two plant communities that made up the true prairie grassland were described above. Detailed description of the ponderosa pine-bunchgrass grass forest followed immediately below.

Students should note the abundant crop of pine cones lying on the ground surface (right foreground) of this grassland.

Powder River County, Montana. Mid-June; late vernal aspect. Mosaic of FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Grama-Needlegrass-Wheatgrass), SRM 606 (Wheatgrass-Bluestem-Needlegrass), Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

11. Inside a forest range- Interior of a ponderosa pine-grass range in Northern Great Plains. This population of ponderosa pine was uneven-aged with all age classes represented. Larger seedlings up to mature trees were visible in these two forest "photoplots". Understorey of this range vegetation was almost exclusively herbaceous with the domi9nant being slender wheatgrass. The main forb was wavyleaf thistle. There were also a few plants of yellow sweet-clover (Melilotus officinalis), an introduced agronomic legume that has naturalized much of the Western Range, especially over most of the Central and Northern Great Plains. Fringed sage (Artemisia frigida) was a suffruticose (sub-shrub) species widespread throughout the understorey of this forest range (see below).

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

12. Understory dominant-Slender wheatgrass ((Agropyron trachycaulum= A. subsecundum= A. caninum) that dominated the herbaceous layer of a ponderosa pine-grass forest range in the Northern Great Plains. Although slender wheatgrass tends to be a pioneer species and one generally adapted to disturbance this short-lived perennial does persist into the climax as part of the understorey of various range types (Tilley et al., 2011). Slender wheatgrass is generally regarded as excellent forage to a number of wildlife and livestock species that is adapted to a number of habitats ranging from wet to relative droughty and from open forest to hillside grasslands (Stubbendieck et al. (1992, p. 205).

Taxonomic note: agrostologists (plant taxonomists in general) have argued for decades over the proper name of slender wheatgrass with first one group of authors (Hitchcock and Chase, 1950; Great Plains "Flora Association, 1986) and then three other groups of authorities (Skinner et al., 1999; Barkworth et al., 2007; Shaw, 2008) arguing for this specific epithet or that genus, each insisting with the arrogance, smugness, and self-rightousness of taxonomic Pharisees that their approach, name, etc. is the only "way, truth, and the light". With advent and ascent of cladistics this "tempest in a teapot" grew even more acrimonious. Rangemen, foresters, agronomists, and horticulturalists are botanical heathen or Gentiles to the high priests of Plant Taxonomy prompting this nonbeliever of such nonsense to offer this, "Damn the lot of them". Call it slender wheatgrass and go on.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

13. Representative specimens- Three different plants of slender wheatgrass in the understorey of an open ponderosa pine forest range in the unglaciated Northern Great Plains. These plants, though of a short-lived perennial species, were part of the climax forest community. Note the needles of ponderosa pine all around.

Powder River County, Montana. Mid-June; immediate pre-anthesis stage.

14. Spiikes beneath the pines- Spike inflorescences of slender wheatgrass produced in an open ponderosa pine forest range in the Northern Great Plains. These spikes which were shown at progressively closer camera (focal) distance were a representative sample of some of those present on plants like the three shown immediately above. The first image included a few of last year's spikes which had shed their spikelets/florets. The rachis of slender wheatgrass dies not disarticulate to the extent of some other members of the Triticeae or Hordeae, wheat or oat tribe.

Powder River County, Montana. Mid-June; immediate pre-anthesis stage.

15. Pine straw and propagules with lower-growing pals- Understorey of ponderosa pine-grass forest range. Ground layer with pine "straw" (shed needles) and pine cones of ponderosa pine and lower herbaceous layer of fringed sage (Artemisia frigida) and some shoots of slender wheatgrass, the dominant of the herbaceous layer of this range plant community.

By whatever name this grass was the dominant--frequently the sole member--of the herbaceous understorey of this forest range.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

Ponderosa pine or western yellow pine is one of the most widely distributed Pinus species in North America, perhaps having the greatest latitudinal distribution of all. Ponderosa pine produces more timber than any other one Pinus species on the continent and it is second only to Douglas-fir in total annual lumber production (Harlow et al., 1979, p. 102). As implied by the common name of western yellow pine, P. ponderosa is in the yellow or hard pine subgenus of Diploxylon. There are three varieties of ponderosa pine: Pacific ponderosa pine (P. ponderosa var. ponderosa), Rocky Mountain ponderosa pine (P. ponderosa var. scopulorum), and Arizona ponderosa pine (P. ponderosa var. arizonica). Students can find details of the silvics of P. ponderosa in such standard texts and references as Harlow et al. (1979, ps.101-106) and Burns and Honkala (1990, Vol. 1, p. 413-424).

Importance of ponderosa pine from perspective of range cover types is as a dominant species that defines cover types, potential natural vegetation (eg. Kuchler mapping units), ecosystems, etc. There are more different forms or variants of understories in ponderosa pine forests than in most other forests or forest cover types, especially those that are defined on basis of dominance. (It was explained in the accompanying review of vegetation units that forest and rangeland cover types are dominance types.) In addition, ponderosa pine is a major (co-dominant to associate) species in forest types made up of several tree species (eg. Black Hills forest covered immediately above, California mixed conifer type, Pacific ponderosa pine-Douglas fir type, gannd fir type).

Within the general or greater region of the southern and central Rocky Mountains there are two ponderosa pine-defined forests that are potential natural vegetation according to the Kuchler system: K-17 (Pine-Douglas-Fir Forest) and K-18 (Arizona Pine Forest). These two Kuchler units plus K-10 (Western Ponderosa Forest), K-15 (Eastern Ponderosa Forest), and K-16 (Black Hills Pine Forest) were covered by Society of American Foresters as Interior Ponderosa Pine (SAF 237). Strictly speaking the ponderosa pine forest in central Arizona is in the Grand Canyon section of the Colorado Plateau physiographic province with specific examples shown from the San Francisco Mountains. The examples of ponderosa pine forest range in central New Mexico and Trans Pecos Texas were in the Sacramento section of the Basin and Range province (Sacramento Mountains and Guadalupe Mountains, respectively). The current publication arranged range-- both forest and rangeland-- cover types by general geographic or natural region and/or the somewhat smaller physiographic provinces within the former. As a result some cover types, especially some forest types, were included under more than one link category or, alternatively, "lumped"-- for convenience-- under a general category that was less specific and less precise. Ponderosa pine was one such example. Taxonomic unit of dominant species was interpreted as more important than physiographic province and section in defining a unit of vegetation (eg. range cover type). Such an arrangement was more consistent with the greater number of vegetation units by the Kuchler potential natural vegetation system than by the Society of American Foresters cover (= dominance) type system.

Kuchler (1964, 1966) distinguished in mapping and description between the Black Hills ponderosa pine and the eastern ponderosa pine forests (units 17 and 16, respectively). In the map accompanying forest and range ecosystems (Garrison et al., 1977) the eastern ponderosa pine forest ecosystem was mapped as unit 15. Examples of this eastern ponderosa pine forest from the Pine Ridge Region and Wildcate Ridge area of western Nebraska (Weaver, 1965, ps. 155-163) were presented next.

16. "East Meets West"- Unique ecotonal forest in Nebraska Sand Hills (also shown as Sandhills) near extreme northern edge (Pine Ridge Escarpment) of High Plains section of Great Plains physiographic province. Weaver (1965, p. 155) described this as stated that the Pine Ridge Escarpment separated the High Plains on the south from the Missouri Plateau (to the north) section of the Great Plains. Range plant communities at such "unions" of landforms are generally "mixtures" of species from adjoining floristic provinces.

The unique vegetation in this photograph is a "three-way" ecotone of tallgrass prairie-eastern deciduous forest-ponderosa pine forest just above the Niobrara River on a bluff-like landform that resulted from wind-blown sand.The grassland community in the foreground was tallgrass prairie of little bluestem, big bluestem, needle-and-thread (Stipa comata), Indiangrass (Sorgastrum nutans), prairie sandreed (Calamovilfa longifolia), and plains lovegrass (Eragrostis trichoides). The largest conifers are of course ponderosa pine, representatives of the easternmost ponderosa pine forest in North America that is a part of the western forest regions. The other tree and shrub species are members of the deciduous forest formation of eastern North America. (Forest range types of this vast formation that are found in Nebraska were included at the end of the chapter designated Southern and Central Forests.) Species in the vegetation portrayed here that are part of the eastern deciduous forest region included eastern red cedar (Juniperus virginiana), shorter conifers at front edge of woody community, bur oak, eastern cottonwood, green ash (Fraxinus pennsylvanica), willow (Salix sp.), paper birch, quaking aspen, American or white elm (Ulmus americana), boxelder (Acer negundo), and eastern hophornbeam (Olstrya virginiana).

Fort Niobrara National Wildlife Refuge, Cherry County, Nebraska. Autumnal aspect, October. "Blend" of FRES No. 39 (Prairie Grassland Ecosytem), K-67 (Nebraska Sand Hills Prairie), SRM 601 (Bluestem Prairie) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). Northwestern Great Plains- Niobrara River Breaks Ecoregion, 43r (Chapman et al., 2001).

17. Landscape and vegetation of Pine Ridge Excarpment and Niobrara River of northern High Plains- Landscape scale view of tallgrass and mixed prairies and eastern ponderosa pine forest vegetation (background) and riparian vegetation (post-shedding of leaves) along Niobrara River (foreground). "Bird's-eye view" of the ecotonal vegetation shown in the preceding slide with tallgrass-mixed prairie grassland transition featured prominently between riparian zone and the eastern ponderosa pine forest in the background. There is also transitional vegetation between ponderosa pine forest and grassland that is a savanna of pines and prairie grasses. (This transition range type was covered as mixed prairie-ponderosa pine savanna under the chapter, Mixed Prairie, Grasslands.)

The small evergreen trees in the center grassland immediately beyond the riparian zone were invading eastern red cedar. Note that the deciduous woody vegetation also lined the draw that extended from the Niobrara River to the downhill edge of the ponderosa pine-dominated community.

Fort Niobrara National Wildlife Refuge, Cherry County, Nebraska. Autumnal aspect, October. "Blend" of FRES No. 39 (Prairie Grassland Ecosystem), K-67 (Nebraska Sand Hills Prairie), SRM 601 (Bluestem Prairie) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). Riparian vegetation was too small to be mapped at scale of these larger vegetation units, but it was certainly no less important. Northwestern Great Plains- Niobrara River Breaks Ecoregion, 43r (Chapman et al., 2001).

A zone of tree-dominated range vegetation developed along the Niobrara River in the Nebraska Sandhills that extended from gallary forest, riparian vegetation, upward (outward from the stream) to a zone of upland forest. Examples of this forest range vegetation were presented immediately below.

Forest vegetation of the Niobrara River Valley contains elements of three general major forest zones: 1) western coniferous forest formation dominated by Pinus ponderosa, 2) eastern deciduous forest formation, and 3) northern boreal forest with paper birch (Betula papyrifera) and quaking aspen X bigtooth aspen hybrid (Populus randidentata X P. tremuloides).

Shrubs, including woody vines, in this general forest community included smooth sumac (Rhus glabra), hop hornbeam (Ostrya virginiana), redosier dogweed (Cornus stolonifer), chokecherry (Prunus virginiana), riverbank grape (Vitis riparia), Virginia creeper (Parthenocissis quinquefolia), thicket creeper (P. vitacea), lanceleaf buckthorn (Rhamnus lanceolata var. glabratus), western snowberry (Symphoricarpos occidentalis), and bittersweet (Celastrus scandens).Major grasses in the riparian vegetation zone presented here included Canada wildrye (Elymus canadensis), marsh muhly (Muhlenbergia racemosa). and the two naturalized Eurasian grasses smooth brome (Bromus inermis) and reed canarygrass (Phalaris arundinacea).

The Niobrara River has one of the most botanically and structurally diverse gallery forest in the Western Range Region. The Niobrara River was designated as the Niobrara National Sceinci River (at least 200 miles of it was) in 1991, but much remains to be done with regard to specific management if it is indeed to be saved and not turned into an irrigation ditch like most of the mid-sized rivers of the North American heartland.

The Niobrara is a braided stream, "a stream that divides into or follows an interlacing or tangled network of several small branching and reuniting shallow channels separated from each other by ephemeral branch islands or cannel bars, resembling in plan the strands of a complex braid" (Wilson and Moore (1998). The definitive work on Niobrara (including geological, biological, and cultural aspects) and was that of Johnsgard (2004).

18. Forest lined- Niobrara River with its floristically rich and structurally diverse gallery forest. There were some open areas of tallgrass prairie, but most of this riverarine range vegetation was comprised of tree-shrub-dominated plant communities. Riverarine vegetation of this gallery forest was descripbed in captions below. Forest Descriptions of the forests and adjoining grasslands along the Niobrara River included Barker and Whitman (1989, p. 18-20), Kantak (1995) and Johnsgard (2004, ps. 45-47), the latter of whom drew much from Kantak (1995).

The hardwood zone of this gallery forest would be of the Elm-Ash or Elm-Ash-Basswood (Plains Haradwood Type) described by Barker and Whitman (1989, p.19).

Cherry County, Nebraska. Late June (estival aspect). Range vegetation descriptions given in following captions.

19. A diverse lot- Gallery forest along the Niobrara River. Vegetation of this gallery forest extended from the riparian zone of sandbar willow (Salix exigua), peachleaf willow (S. amygdaloides), red osier dogwood (Cornus stolonifera), and some chokecherry (Prunus virginiana) through a middle zone of green ash (Fraxinus pennsylvanica), American elm (Ulmus americana), box elder (Acer negundo), basswood or American linden (Tilia americana), eastern cottonwood, and bur oak (Quercus macrocarpa) and hackberry (Celtis occidentalis) to the highest and outermost forest zone dominated by ponderosa pine and eastern red cedar (Juniperus virginiana). Obviously there was considerable overlap of many of these species especially in the broad mid-zone to lower parts of the outermost forest edge. Shrubs (in addition to those listed for the riparian zone) included riverbank grape (Vitis riparia), Virginia creeper (Parthenocissis quinquefolia), thicket creeper (P. vitacea) and poison ivy (Toxicodendron radicans= Rhus radicans ).

There were very few forbs in this dense forest, at least at this late spring-early summer season. Most grass was along the moist edge of the bank and consisted of naturalized mooth brome. There was some bottomland switchgrass and Canada wildrye. Also some caric sedges (Carex spp.).

Cherry County, Nebraska. Late June (estival aspect). FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest) for the central hardwood zone while the lower or riparian zone could be interpreted as K-89, Northern Flood Plain (Populus-Salix-Ulmus). SAF 235 (Cottonwood-Willow) for riparain zone; SAF 93 Sugarberry (Hackberry variant)-American Elm-Green Ash for central zone; SAF 237 (Interior Ponderosa Pine) for driest, upper zone.Not a relevant unit for most of this forest in Brown et al. (1998).Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

20. Braided and lined- This view spanning the Niobrara River showed the stream channel feature of this braided stream. Braided streams have usually been interpreted as having loads of sediment that exceed their capacity to carry them (Wilson and Moore, 1998). The result is deposition of these overloads in the network of small, shallow interchannels and channel bars. Range vegetation along this stretch of the Niobrara River was described in the next two sets of slides.

Cherry County, Nebraska. Late June.

21. Farthest edge of sandhill gallery forest- At uppermost (least mesic) zone of the gallery forest hat developed along the Niobrara River ponderosa pine and easter red cedar co-cominated the vegetation. This was where the forest contacted tallgrass prairie of upland switchgrass, big bluestem, little bluestem, sideoats grama, and blue grama with some cheatgrass. The most abundant (about the only) shrub was smooth sumac (Rhus glabra). Ponderosa pine had invaded the grassland here at the dege of these two types of range vegetation. This may have been due to greater tolerance of this species to fire.

Cherry County, Nebraska. Late June. Forest community was 21 (Pondaerosa Pine Forest and Woodland Ecosystem). K-15 (Eastern Ponderosa Pine Forest with Eastern Red Cedar do-dominant). SAF 237 (Interior Ponderosa Pine) In Brown et al. (1998, p. 37): eastern form of Yellow Pine Series, 122. 62) Grassland was FRES No. 39 (Prairie). K-67 (Nebraska Sand Hills Praire) or, perhaps, more precisely at this location, K-66 (Bluestem Prairie). Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

22. Forest meets prairie in the Nebraska Sandhills-Panarama of uppermost zone of gallery forest above Niobrara River coming into contact with tallgrass prairie. At higher (and drier) portions of this riverarine forest some scattered boxelder and bur oak joined co-dominant ponderosa pine and eastern red cedar to make a "last stand" at edge of the regional climax of tallgrass prairie. The prairie was dominated by upland switchgrass with big bluestem, little bluestem, and Canada wildrye associated major species.

Cherry County, Nebraska. Late June. Forest community was 21 (Pondaerosa Pine Forest and Woodland Ecosystem). K-15 (Eastern Ponderosa Pine Forest with Eastern Red Cedar do-dominant). SAF 237 (Interior Ponderosa Pine) In Brown et al. (1998, p. 37): eastern form of Yellow Pine Series, 122. 62) Grassland was FRES No. 39 (Prairie). K-67 (Nebraska Sand Hills Praire) or, perhaps, more precisely at this location, K-66 (Bluestem Prairie). Nebraska Sandhills-Sand Hills Ecoregion 44a (Chapman et al., 2001).

23. Eastern (Pine Ridge Escarpment) ponderosa pine forest- Interior of the open-- almost woodland-like physiogonomy-- form of ponderosa pine on the Great Plains at edge of Nebraska Sand Hills prairie. This vegetation could be interpreted as a mixed prairie grassland-western yellow pine forest transition (ie. a savanna of ponderosa pine and mixed or, even, tallgrass prairie; ecotonal vegetation between eastern edge of the western coniferous forest region and Great Plains grasslands). Equally valid in the mind of this author was the interpretation of regenerating ponderosa pine forest of the Pine Ridge Region as described by Weaver (1965, ps. 155-161). Weaver (1965, ps. 160-161) included the deciduous tree species of riparian areas and canyons as part of the regional evergreen forest of the Pine Ridge.

Soapweed yucca (Yucca glauca) growing atop the ridge in foreground and little bluestem (the brown bunchgrass) growing alongside the yucca and beneath pines were indicators of the Sand Hills prairie element that comprised the understorey of this vegetation.

This range vegetation was included as an example of the eastern form of ponderosa pine forest. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-15 (Eastern Ponderosa Forest). SAF 237 (Interior Ponderosa Pine). Fort Niobrara National Wildlife Refuge, Cherry County, Nebraska. Autumnal aspect, October.

Note: Background knowledge of the portion of the Great Plains province on which this range vegetation developed was in the definitive authority of Fenneman (1931, ps.17-21) and the exceptional bulletin by Trimble (1990, ps. 29-32). Northwestern Great Plains- Niobrara River Breaks Ecoregion, 43r (Chapman et al., 2001).

Pine Ridge Region ponderosa pine-Within the Pine Ridge Region ponderosa pine grows as open understorey forests, woodlands, or savannahs with distinctions being cover and density of pines. In theory of Landscape Ecology these forests, woodlands, or savannas were patches within a matrix of mixed prairie or, in case of the escarpment above the Niobrara River, tallgrass prairie.

Ponderosa pine-dominated or -defined range plant communities that had developed within the general mixed prairie association had varying dominat and associate species. Dominants typically included needle-and-thread (Stipa comata), blue grama (Bouteloua gracilis), Junegrass (Koleria cristata= K. pyrimidata), and western wheatgrass. Other locally important native grasses varied from tallgrass species like little bluestem (Andropogon scoparius), big bluestem (A. gerardii), prairie sandreed (Calamovilfa longilfolia) through midgrasses including sideoats grama (B. curtipendula) and Canada wildrye (Elymus canadensis) through to shortgrass such as buffalograss (Buchloe dactyloides), Sandberg's bluegrass (Poa secunda), plains bluegrass (P. arida), and hairy grama (B. hirsuta). The native annual sixweeks fescue (Festuca octoflora= Vulpia octoflora) was frequently well-represented. Annual weedy Eurasian grasses, notably cheatgrass (Bromus tectorum) and Japanese brome or chess (B. japonicus), were common on ranges degraded by disturbances like overgrazing or out-of-proper-season fires. Naturalized perennial grasses included smooth brome (B. inermis) and Kentucky bluegrass (P. pratensis). A locally common, even dominant grasslike speceies was threadleaf sedge (Carex filifolia). Generally the most common forb was fringed sagewort (Artemisia frigida). Otherwise-and surprising to this author-was the conspicuous scarcity of composites. Native legumes included silvery lupine (Lupinus argenteus) and Missouri milkvetch (Astragalus missouriensis). Scarlet globemallow (Sphaeralcea coccinea) had an infrequent though colorful presence. Shrubs other than soapweed (Yucca glauca) and local stands of chokecherry (Prunus virginiana) were basically absent from these open understorey ponderosa pine communities.

Concise yet classic coverage of the forest range vegetation of the Pine Ridge district was given by Weaver (1965, psw. 155-161).

24. Pines in the prairie- An island of eastern ponderosa pine in the vast mixed prairie region of the Central Great Plains. In this semiarid zone there are portions of the Pine Ridge district on which open forest or, in some cases, even more open woodlands dominated by ponderosa pine with a grassy herbaceous layer developed. These climax range plant communities amount to pines growing on a mixed prairie of tallgrass, midgrass, and even some shortgrass species. Sometimes eastern red cedar is a distant associate tree species in this cover type although such was not the case in this example

Grasses under pines were essentially the same as those on adjacent mixed prairie. Dominant species were needle-and-thread, bule grama, and western wheatgrass. At this early summer stage the cool-season needle-and-thread had substantially more biomass and cover than the warm-season blue grama. Following a wet yet cold spring, western wheatgrass was abnormally limited in its cover and herbage yield whereas the current temperature-soil moisture combination resulted in unusually favorable growing conditions for needle-and-thread. Junegrass was a locally common grass. Less common grasses as well as forbs were listed in the introduction to this sample of Pine Ridge ponderosa pine range.

Organization note: This mixed prairie range type, SRM 608 (Wheatgrass-Grama-Needlegrass), was treated in the grassland chapter, Mixed Prairie. It was not described or discussed at this juncture.

Sioux County, Nebraska. Late June (estival aspect). Ponderosa pine forest range community was FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-15 (Eastern Ponderosa Forest). SAF 237 (Interior Ponderosa Pine). Rocky Mountain Montane Conifer Forest 122.6,Yellow Pine Series 122.62, Pinus ponderosa Association 122.621 (Brown et al., 1998, p. 37). Western High Plains-Pine Ridge Escarpment Ecosystem, 25a.

25. Pine-clad draw- Along and inside of a draw, ravine, or small canyon ponderosa pine of various age/size classes formed an arboreal component above a mixed prairie to produce an open canopy ponderosa pine forest with a grassy understorey. Both of these photographs were taken at points of contact of this forest with a mid-grass prairie. Dominant grasses in both range plant communities were needlt-and-thread, blue grama, western wheatgrass with associate and incidental grasses including Junegrass along with hairy grama, big bluestem, Sandbeerg's bluegrass, little bluestem, smooth brome, cheatgrass, Japanese brome, and sixweeks fescue. Threadleaf sedge was common in some spots. There were relatively few forbs and almost no shrubs. The most common forb was fringed sagewort. Composition of the sward varied locally depending on slope, shade, disturbance (eg. erosion), etc.

Sioux County, Nebraska. Late June (estival aspect). FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-15 (Eastern Ponderosa Forest). SAF 237 (Interior Ponderosa Pine). Rocky Mountain Montane Conifer Forest 122.6,Yellow Pine Series 122.62, Pinus ponderosa Association 122.621 (Brown et al., 1998, p. 37). Western High Plains-Pine Ridge Escarpment Ecosystem, 25a.

26. Three years after the flames- Panaramic views of a Pine Ridge ponderosa pine-dominated plant community that ranged from an open forest to a woodland (tree crowns typically not in contact with each other) that had been burnt by a crown fire about two years earlier. This forest range vegetation was now in its third growing season following the intense fire. There had been no livestock grazing. Influence of deer feeding was not known.

This forest had not been logged nor treated with prescribed burning. The fire occurred in summer (ignition by dry lightening during a thunderstorm) of the second or third year of a worse-than-typical drought. Fuel-loading was an on-going process in absence of proper fuel management (ie. no fuel-reduction prescribed fires because Smokey's friends "prevent forest fires"). Everything was in place for the "holocaust" that ensued--as it always does sooner or later with such mismanagement or negligence, justified by "not playing with matches". The confragation did furnish a unique opportunity to document and analyze natural reforestation and secondary plant succession in this forest cover type.

In addition to remaining ponderosa pine (the defining dominant species) other major plants (ie. those with greatest cover, density, biomass) on these burntover hills were smooth bromegrass and cheatgrass, both naturalized cool-season Eurasian species. The reddish spots were primarily cheatgrass with some Japanese brome or Japanese chess. Most ("far and away" the greatest) cover in green areas was smooth brome, the most commonly seeded agronomic forage grass in this region, which had naturally invaded the scorched soil surface of this former climax forest range vegetation.

Native grasses, a "natural blend" of both warm- and cool-season perennials, were starting to re-establish, but at this time they had been overwhelmed by smooth brome. Dominant native grasses were needle-and-thread, blue grama, western wheatgrass with Junegrass, big bluestem, little bluestem, sideoats grama, Sandberg's bluegrass, and, more locally, prairie sandreed, buffalograss, and hairy grama. Kentucky bluegrass, another introduced pasture grass, was also present, though under these conditions of no livetock grazing, this shorter-growing species was not competitive with smooth brome (or the annual Mediterranean bromes).

Chokecherry was an infrequent shrub other than locally (eg. lower left corner in first of these slides).

Ponderosa pine seedlings had begun to re-stock this fire-demolished Pine Ridge ponderosa pine forest (subject of next photograph).

Sioux County, Nebraska. Late June (estival aspect). FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-15 (Eastern Ponderosa Forest). SAF 237 (Interior Ponderosa Pine). Rocky Mountain Montane Conifer Forest 122.6,Yellow Pine Series 122.62, Pinus ponderosa Association 122.621 (Brown et al., 1998, p. 37). Western High Plains-Pine Ridge Escarpment Ecosystem, 25a.

27. Up from the ashes- Successional recovery of vegetation on a Pine Ridge ponderosa pine-dominated open forest (in some areas more of a woodland) in the third growing season following a crown fire that drastically disturbed this native range plant community. Two- or, at most, three-year-old seedlings of ponderosa pine wer regrowing on the "remains" of this partially denuded (largely "de-treed") forest on the semiarid Nebraska Pine Ridge Excarpment.

The major grass in this view of recovering forest was smooth brome, a species that had naturalized in this region following its introduction as a domestic forage crop. On the south slope in the background the reddish patches were cheatgrass, a naturalized Mediterranean annual. Native tall- and midgrass species (listed in the above introduction) had made notable recovery, but were still lagging behind the exotic species which are well-adapted to disturbance.

This was a natural, ready-made experiment to document recovery of forest vegetation.

Sioux County, Nebraska. Late June (estival aspect). FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-15 (Eastern Ponderosa Forest). SAF 237 (Interior Ponderosa Pine). Rocky Mountain Montane Conifer Forest 122.6,Yellow Pine Series 122.62, Pinus ponderosa Association 122.621 (Brown et al., 1998, p. 37). Western High Plains-Pine Ridge Escarpment Ecosystem, 25a.

29. Southwestern Ponderosa Pine Forest- Texas is so big and and situated such as to be composed of seven (that is correct, 7) of the 29 biotic provinces of North America (Dice,  1943). These extend from the Austroriparian which includes the Pineywoods forests of east Texas through the Chihuhuan of the Trans Pecos Basin and Range physiographic province. Dice (1943, p. 41) extended a small portion of the Navahonian biotic province into the Guadalupe Mountains of the Trans Pecos region. Turner (1959, ps. 6-7) discussed the biotic province scheme of Dice but mapped the northern part, including the Guadalupe and neighboring ranges, of the Trans Pecos region as in the Rocky Mountain floristic province. In the Guadalupes there is a montane zone dominated by ponderosa pine with Douglas-fir on north slopes.

Note: Kuchler unit 17 included all ponderosa pine types in the southwest and the central Rocky Mountains whereas Kuchler separated out other ponderosa pine types such as Western Ponderosa Forest (K-10), Eastern Ponderosa Forest (15), Black Hills Forest (K-16), and Arizona Pine Forest (K-18). Obviously this is valid using overstory features (among others). It is also valid, however, to distinguish the ponderosa pine range types of the southern Rockys from those of the Colorado Rockys (to distinguish within K-17) on basis of different dominant grasses, forbs, and shrubs in the lower layers of understory.

31. A little bit of everything- Ponderosa pine-mixed shrub-montane grass savanna in the upper Front Range of the Southern Rocky Mountains. Lower montane conifer forest of Dick-Peddie (1993, p. 66), but none of the series in New Mexico were germane this far north. This was the Ponderosa Pine Ecosystem of Beidleman et al. (2000, p. 11) with thimbleberry (Rubus parviflora) and wax currant (Ribes cereum) as main (=co-dominant) shrubs--at least locally--and antelope bitterbrush (Purshia tridentata) as the associate shrub. Numerous plants of thimbleberry and wax current were present in foreground of these slides.

Major grass species were blue grama (Boutelous gracilis) and an assortment of Poa species including Kentucky bluegrass (P. pratensis), Sandberg's bluegrass (P. secunda= P. sandbergiii), and Wheeler's bluegrass (P. wheeleri= P. nervosa) along with various bromes including fringed or mountain brome (Bromus ciliatus and/or B. richardsonii) and nodding brome (B. anomalus= B. porteri) plus mountain muhly (Muhlembergia montana) and probably other muhlies as well. There were also several caric sedges (Carex spp.) present.

The most abundant forb on this ponderosa pine-shrub-grass savanna at this relatively early stage of the growing season was mountain goldenbanner or mountain goldenpea (Thermopsis montana= at least in part, T. rhombifolia). Another conspicuous and locally abundant forb was Rocky Mountain locoweed, white locoweed, or silky crazyweed (Oxytropis sericea). Both of these papilionaceous legumes are toxic--under certain conditions--to range animals.

Warning (especially to traditionalists): *$#**+ taxonomists changed binomials of many of these species such as thimbleberry to Rubacer parviflorum (Weber, 1990) and, in the ultimate blasphemy, blue grama to Chondrosum gracile (Shaw, 2008)!

This was an example of ponderosa pine savanna with both shrubs and herbaceous species locally dominant (ie. a mosaic form of ponderosa pine range.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect). FRES No. 21 (Ponderosa Pine Ecosystem). Mapped as Kuchler- 17 (Ponderosa Pine-Douglas-fir Forest). SAF 237 (Interior Ponderosa Pine); no SRM for this pine type per se, but it would be the Southern Rocky Mountain equivalent of SRM 109 or 109 and 110 combination (Ponderosa Pine-Shrubland-Grassland). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

32. Hale and hearty specimen- Fine (and seemed to be quite old) specimen of thimbleberry (Rubus parviflorus) growing on an open ponderosa pine montane range on Front Range of Southern Rocky Mountains. Thimbleberry is an extremely widely distributed species ranging from Alaska south to British Columbia and east to Manitoba and south into northern Mexico. It occurs in all eleven of the westernmost (continental) Range States.

As a browse plant thimbleberry was reported (Dayton, 1931, p. 58) to be "mostly poor or worthless" though it was infrequently utilized lightly by cattle. In the Range Plant Handbood (Forest Service,937, B139) it was concluded that palatability of thimbleberry "varies considerably in different localities" varying from poor to fair for cattle and fair to good for sheep. Similarily, feed value of thimbleberry varied for the cervids. Utilization of thimbleberry was found to be heavy when other (and more palatable) browse was of limited availability.

Associated shrubs on the example of ponderosa pine montane range shown here included antelope bitterbrush (Purshia tridentata) and wax currant (Ribes cereum).

Rocky Mountain National Park, Larimer County, Colorado. Mid-June.

34. Another hearty specimen (of another species)- A large plant of wax currant (Ribes cereum) growing on the open ponderosa pine montane range (along Front Range of Southern Rocky Mountains) featured here. This plant was a neighbor of the large thimbleberry just presented. There was also antelope bitterbrush in the shrub layer of this foresst range.

Dayton (1931, p. 41) and the Forest Service (1937, B131) concluded that although wax currant was not particularily palatable to grazing animals the abundance of this species and the browse it offered was so great that this species was important in grazing capacity of ranges on which it grew. Reportedly, wax currant browse was found to have "high protein content". Also, absence of spines on wax currant combined with openness of grazing lands on which this shrub occurred made for beneficial utilization of this browse plant.

It was interesting--though revealing--that the best and most practical discussions (brief though they were) of wax current and thimbleberry as range plants were those that had been written decades ago.

Mountain National Park, Larimer County, Colorado. Mid-June, full-bloom stage.

35. Leader of wax current- Terminal portion of shoot of Ribes cereum showing leaves and inflorescences of one of the wide-ranging and important shrubs in the Western Range Region.

Mountain National Park, Larimer County, Colorado. Mid-June, full-bloom stage.

36. Rocky Mountain gold (and poison)- Mountain goldenbanner or mountain goldenpea (Thermopsis montana= at least in part, T. rhombifolia) growing on a ponderosa pine savanna in the foothills of the Front Range. This very showy plant is also a poisonous one, yet one with a confused nomenclatural past such that there could be something of a case of "mistaken identity". Burrows and Tyrl (2003, ps. 617-620) described toxicity attributed to three Thermopsis species, one of which on western ranges was T. rhombifolia but which was often reported as T. montana. T. rhombifolia is now recognized as being "highly polymorphic" so that a number of forms previously interpreted as separate species ae now included in an expanded T. rhombifolia (Burrows and Tyrl, 2003, p. 618). Weber (1990, p. 195) clearly distinguished between T. montana. and T. rhombifolia specifying that in addition to morphological features the former grew in mountain meadows and parks whereas the latter was "strictly a plains species".

Anyway, mountain goldenbanner was a striking range legume on the open ponderosa pine savanna and open forest ranges. T. montana was featured in Notes on Western Range Forbs (Hermann, 1966, ps. 136-137) in which it was stated the this "is by far the commonest western species, as well as the most variable". Apparently its palatability also "varies enormously". Previously this species was included in the Range Plant Handbook (Forest Service, 1937, W186) were it was reported that it was grazed mostly under "severe overgrazing". Goldenpeas "often increase on ranges which have been overgrazed". "Goldenpeas, practically worthless as forage, are important as range plants because of their wide distribution and commonness."

Rocky Mountain National Park, Larimer County, Colorado. Mid-June.

37. Beward of banner- Inflorescence of papilionaceous flowers of mountain goldenbanner. This is one of the more conspicuous range forbs of ponderosa pine-dominated savanna, woodland, and open forest ranges.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June.

38. Gone crazy- Rocky Mountain locoweed, white locoweed, or silky crazyweed (Oxytropis sericea) on an ecotone between ponderosa pine open forest and quaking aspen (Populus tremuloides) parkland. This is one of the "classiest" of the poisonous range plants: "classy" as in showy or stuningly attractive and as a classic or textbook poisonous range plant. Silky crazyweed is one of several Oxytropis and Astragalus species regarded by Kingsbury (1964, ps. 307, 309) as "true locoweeds" (ie. produce typical symptoms of loco poisoning). This species usually made it into the standard texts of poisonous plants (eg. Cheeke and Shull, 1985, ps. 143, 146). In their poisonous plant compendium Burrows and Tyrl (2003, ps. 594-599) covered Oxytropis species, including O. sericea. (Incidentially this magnificant reference is highly recommended to all true rangemen. It should be a shelf companion to Kingsbury [1964].)

Hermann (1966, ps. 124-125) remarked that O. sericea was almost as important as a stock-poisoning plant as the better-known O. lambertii , Lambert's crazyweed, except that livestock are less apt to feed on O. sericea.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June, and can there be any debate that it was at full-bloom stage?

40.  Ponderosa pine forest range typical of the Colorado Rockies. Pine dropseed in full bloom and dominating the herbaceous understory shows the potential forage yields of one form or phase of this very widespread forest range type.Scattered Douglas fir and a shrub layer dominated by mountain mahogany and wax current illustrate the botanical diversity of even a park-like western coniferous forest.

Rocky Mountain National Park, Colorado. August. FRES No. 21 (Ponderosa Pine Ecosystem). K-17 (Ponderosa Pine- Douglas-fir Forest). SAF 237 (Interior Ponderosa Pine). No SRM description for the ponderosa pine cover type in the central or southern Rocky Mountains. Pinus ponderosa-mixed conifer Association in Yellow Pine Series of Brown et al. (1998). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

41. Big natural park in Southern Rockies- Landscape-scale panarama of a mountain grassland of Arizona fescue (Festuca arizonica) and mountain muhly (Muhlenbergia montana) within a surrounding forest of ponderosa or western yellow pine in the Southern Rocky Mountains. This mountain grassland occupied a nearly treeless valley having an area of approximately five square miles that was situated within mountain slopes covered with ponderosa pine forests, sometimes with Engelmann spruce (Picea engelmannii) as the associate tree. In the bottom of this valley range plant communities were primarily montane marshes usually called wet subalpine mountain meadows. Between the wet meadows and this Arizona fescue-Mountain muhly steppe (bunchgrass prairie) there were vegetational zones of moist to dry meadow (dry subalpine mountain meadow). The range vegetation of meadows varied considerably, but consisted mostly of caric sedges, spikerushes, rushes, and cool-season grasses (especially Poa species). The term applied collectively to these herbaceous plant communities is park.

Park is the term or designation applied to such areas of herbaceous vegetation that exist as "grassy openings" or glades within a larger vegetation or plant community that is more-or-less forest or woodland. Parks can be extremely large so as to encompass thousands of acres as for example North, Middle, and South Parks in the Colorado Rockies or quite small as, say, less than ten acres (maybe less than one acre). Other parks can be of intermediate size that are still relatively large such as the one shown here. Park apparently originated as part of the local idiom among rural folk tracing pack to mountain men and miners. The word become widely accepted as a term applied with situationally dependent meanings and connotations including as a designation for natural vegetation. Park even has cultural usage such as to distinguish political units in, for instance, Park County, Colorado where range vegetation shown in this and several succeeding photographs had developed.

The range vegetation featured here was mountain steppe dominated by Arizona fesuce (with some Idaho fescue [Festuca idahoensis]) and mountain muhly (with some slimstem muhly [Muhlembergia filiculmis]). In some restricted locations there was a savanna to woodland of ponderosa pine as a tree layer to this montane grassland. (See next slide). Grasses were limited to these species as dominants and associates with scattered plants of various needlegrasses several of which either integrade or are very similar. The latter included Letterman's needlegrass (Stipa lettermannii) which, according to various authorities, has frequently included--wholly or in part--Nelson's needlegrass (S. nelsonii), pine needlegrass (S. pinetorum), and/or western neddlegrass (S. occidentalis). Search as he might the author did not find any cheatgrass (or any other species of annual grass) nor bluegrass species in this grassland of cespitose grasses. .

San Isabel National Forest, Park County, Colorado. Mid-June (late vernal aspect). Mosaic of: FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine) and FRES No. 36 (Mountain Grasslands Ecosystem), K-46 (Fescue-Mountain Muhly Prairie), no SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

42. Pine in big natural park- Local savannah or woodland of ponderosa pine on part of an Arizona fescue-mountain muhly steppe in a large park in the Southern Rocky Mountains. This slide showed a portion of the Arizona fescue-mountain muhly montane grassland (a bunchgrass park) introduced in the preceding photograph from a slightly different location to give an example of local grove-like stands of ponderosa pine on what was otherwise a mountain grassland made up of only a few cespitose grass species. Other grasses in this range plant community included Idaho fescue, slimstem muhly, and Letterman's needlegrass in about that order of abundance (which was not abundant at all).

In this general location there was a continuum of natural vegetation ranging from fescue-muhly steppe (grassland of cespitose or tufted species of two Gramineae genera), savanna as a fescue-muhly community with widely scattered ponderosa pine, woodland of ponderosa pines that did not contact each other over a prominent fescue-muhly layer, and finally to ponderosa pine forests with tree densities that permitted only limited development of an herbaceous understorey. This different local plant communities occurred as a mosaic. This pattern of range vegetation was shown in the next photograph below.

San Isabel National Forest, Park County, Colorado. Mid-June (late vernal aspect). Mosaic of: FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine) and FRES No. 36 (Mountain Grasslands Ecosystem), K-46 (Fescue-Mountain Muhly Prairie), no SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

43. A studded landscape- A landscape-scale view showing a mosaic of montane grassland and forest in the Southern Rocky Mountains at an elevation of roughly 9000 feet. This range vegetation was at slightly higher elevation than the larger park described in the two immediately preceding photographs. This grassland was also a mountain steppe (bunchgrass prairie), but forests while were primarily ponderosa pine also included Engelmann spruce as the associate to local dominant. In the perspective of Landscape Ecology these forests were patches in a matrix of montane prairie made up of cespitose (tufted or clumped) grasses the major species of which were Arizona fescue and mountain muhly. In addition to these dominants associate grass species included Idaho fescue, slimstem muhly, Nelson's needlegrass, and glaucus bluegrass (Poa glauca). There few forbs present were mostly composites and of no apparent consequence at this season (in this plant society).

At larger (area-wide or mountain range) scale montane grasslands were patches within a forest matrix. In other words, there were smaller-size vegetational units that could be seen as patches immersed within larger-size vegetational areas that could be viewed as patches within larger-yet units of vegetation which could be regarded as the general vegetation that qualified as the all-encompassing matrix. Contrary to early anticipation of the newer discipline of Landscape Ecology, landscape theory and perspective did not eliminate the subjectivity of the earlier paradigm of Ecosystem Ecology that often posed problems in delinating boundaries of ecosystems. All ecological units whether natural plant-animal communities (biomes, associations, etc.), ecosystems, landscapes, populations, species, or even individuals of clonal organisms pose the same problem as to "where to draw the line".

But to return to this range vegetation: the local tree-dominated communities were mostly stands consisting of about five or six up to twenty (at the most) ponderosa pine and some Engelmann spruce with most conifers being of mature age and size with all together forming in tiny to small forest clumps within an otherwise uninterupted mountain steppe. Engelmann spruce tended to be larger than the ponderosa pine. For comparison, the two largest trees in right-center midground were Engelmann spruce which, incidentially, had produced an abundant cone crop.

The larger, taller, and more-rank tufts of grass were Arizona fescue, the local dominant. Grass clumps of shorter height were mostly montain muhly, the local associate species, with some Idaho fescue and slimstem muhly. These grasses are strictly cespitose species meaning that they are tufted with all their shoots being tillers (no horizontal shoots such as rhizomes). Thus this grassland physiogonomy is that of a steppe or bunchgrass prairie.

San Isabel National Forest, Park County, Colorado. Mid-June (late vernal aspect; all grasses in preboot phenological stage). Grassland vegetation was FRES No. 36 (Mountain Grasslands Ecosystem), K-46 (Fescue-Mountain Muhly Prairie), no SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Forest community (patches or clumps thereof) was FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

44. High elevation grassland with some tree neighbors- A montane grassland of cespitose grasses that formed a high-elevation (about 9000 feet) steppe (bunchgrass prairie) dominated by Arizona fescue and with mountain muhly as associate (in a few local habitats these were co-dominant species). In certain locations Arizona fescue grew in consociations of relatively extensive coverage. The second of these two photographs presented an example of a consociation or single-species stand. Other--though always minor--grasses included Idaho fescue, slimstem muhly, Letterman's needlegrass, and even glaucus bluegrass. Cheatgrass had not invaded this grassland. There were no forbs of apparent importance at this season (in this vernal plant society).

Two conifer species grew in aggregations that formed clumps of five or six up to roughly twenty adult (though relatively modest-sized) trees. The dominant of these was western yellow or ponderosa pine (all trees in the first slide). Engelmann spruce was the associate conifer based on relative cover and density. The largest trees (height and trunk diameter), however, were Engelmann spruce (eg. the two isolated trees at right-of-center skyline in the second slide). Although this elevational zone was below timber (tree) line, the Engelmann spruce showed some general features caused by wind-pruning. An example of this was the numerous shrublike trees and/or shoots of Engelmann spruce that had formed at the base of the two adult trees just mentioned.

Any close spatial association among woody plants and herbaceous ones raises questions regarding competition and survival in the presence of each other. Obviously bunchgrasses and ponderosa pine have co-existed for millennia, but there could still be other than mutually beneficial symbiosis. Larson and Lachubert (1969) reported that roots of Arizona fescue and mountain muhly grew faster than roots of ponderosa pine suggesting potential competition between ponderosa and its two major understorey herbaceous species. In fact, Rietveld (1975) found that competition from these two grasses could suppress or even kill seedlings of ponderosa pine. Occurrence/presence of trees either in small groups (like "mini- forests"), as here, or as woodlands or savanna, as shown above, suggested that for some reason(s) co-existence of these species of conifers and bunchgrasses was limited. Fire history of this range vegetation was unknown to the author.

Park County, Colorado. Mid-June (late vernal aspect; all grasses in preboot phenological stage). Grassland vegetation was FRES No. 36 (Mountain Grasslands Ecosystem), K-46 (Fescue-Mountain Muhly Prairie), no SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Forest community (patches or clumps thereof) was FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

45. Mountain steppe- Physiogonomy, structure, and species composition of montane bunchgrass prairie with Arizona fescue and mountain muhly as dominant and associate, respectively, or co-dominants locally. There were some plants of Idaho fescue, slimstem muhly, Nelson's needlegrass, and even glaucus bluegrass although all of these were present only as scattered to widely scattered individuals. An unusually dense--though still local--group of glaucus bluegrass plants was in the center foreground of the second slide. These plants were in early stages of grain production. Gaucus bluegrass was the only grass even approaching flowering. Arizona fescue is a later-maturing cool-season species while mountain muhly is a warm-season, C4 grass (Shaw, 2008, ps. 157, 399). Cheatgrass was not present nor were there forbs of obvious ecological importance at this season. The few forbs present were composites.

The second photograph presented greater detail of the interrupted sward of this mountain bunchgrass prairie.

Range vegetation seen in these two slides was on the same grassland as shown in the three immediately preceding slides.

San Isabel National Forest, Park County, Colorado. Mid-June (late vernal aspect; all grasses in preboot phenological stage). FRES No. 36 (Mountain Grasslands Ecosystem), K-46 (Fescue-Mountain Muhly Prairie), no SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

46. Ponderosa pine-bunchgrass forest range- Three overall views of the open form of a montane forest (or forestlike) community with the broken (interrupted) tree canopy layer dominated by ponderosa pine and one or two herbaceous layers dominated by grasses, especially cespitose species, in the following, approximate relative order (as found on the range presented here at time of photographs): mountain muhly, Arizona fescue, pine dropseed, Indian ricegraass, bearded or slender wheatgrass (Agropyron subsecundum= A. trachycaulum), New Mexico sleepygrass (Stipa robusta), Nelson's needlegrass (S. nelsonii), purple reedgerass or purple pinegrass (Calamagrostis purpurascens), blue grama, muttongrass or Fendler's bluegrass (Poa fendleriana), and bottlebrush squirreltail (Sitanion hystrix). Sun caric sedge (Carex heliophila= C. inops subsp. heliophila= C. pensylvanica subsp. helipophila= C. exiebenia) was the most common or abundant grasslike plant. At the early summer season when these photographs were taken sun sedge had considerably greater visible density and cover than minor grass species (on the range shown here) like blue grama, mutton bluegrass, and bottlebrush squirreltail.

Forbs were quite restricted or widely spaced but included a number of species. Forb species readily identified at time of photographing included lodge lupine (Lupinus parviflorus), wholeleaf Indian paintbrush (Castilleja integra), one-sided penstemon (Penstemon unilteralis), Rocky mountain pussytoes (Antennaria parvifolia), sagewort wormwood or field sagewort (Artemisia campestris), Fremont's geranium (Geranium caespitosum fremontii), Fendler's groundsel or Fendler's ragwort (Senecio fendleri), Lambert's crazyweed (Oxytropis lambertii), miner's candle (Cryptantha virgata), and prairie thistle (Cirsium canescens).

Shrurbs were even more limited (again, on the range viewed here). The most common shrub was either Arkansas rose (Rosa arkansana) followed by wax current (Ribes cereum) and skunkbush sumac (Rhus trilobata= R. odorata).

A handy species list and fine summary of research on this forest range was that prepared by Gary (1985).

This ponderosa pine-bunchgrass range community was more technically a woodland given that crowns of mature pines did not interlock as generally specified to distinguish froest from woodland. Much of the area was strictly specking even more of a savanna as can be clearly seen from in some of the slides (see those below in particular). The woodland/savanna feature was recognized in classification of this natural vegetation by the Colorado Natural Heritage Program as shown below.

The herbaceous understorey in the first two of these three slides was dominated overwhelmingly by montain muhly while the herbaceous layer in the forest range seeen in the third slide was a consociation of slender or bearded wheatgrass. The limited number of young trees in this ponderosa pine woodland was shown in the first photograph were a few pines of sapling and pole sizes were visible. More examples of limited regeneration were shown below at shorter focal distances.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

47. Typifying the ponderosa pine-bunchgrass type- Composite views of structure and composition of an open (interrupted or sporadic) canopy, low-density (wide tree spacing) ponderosa pine forest (woodland is the more exact designation) with a intermediate-height (midgrass) understorey. This herbaceous component was primarily one layer although in local areas it had herbaceous species of varying heights (eg. some New Mexico sleepygrass of heights in excess of five feet, Rocky Mountain pussytoes of less than six inch height, typical midgrasses like blue grama and pind dropseed) resulting in a "staggered" or fluctuating height (or depth) of the lower layer of vascular plants.

In most parts of this montane coniferous woodland the herbaceous zone was a consociation of mountain muhly. In fact, this cespitose eragrostoid grass formed exclusive stands so that the herbaceous layer was a population of this species and not an herbaceous community or mixed species layer. Other grasses as well as forbs and shrubs were listed in the immediately preceding caption. In these two "photoplots" the only other grass species of ecological or managerial consequence were a few widely scattered plants of New Mexico sleepygrass and slender or bearded wheatgrass. Arizona fescue (Festuca arizonica) occurrs in ponderosa pine woodlands in the area of which the range vegetation shown here was part, but this Arizona fescue was (could) not be located and/or identified from tattered shoots by this rangeman. Arizona fescue has often been described as being co-dominant with mountain muhly in the ponderosa pine-bunchgrass ranges of the Southern Rocky Mountains, including on parts of the Manitou Experimental Forest (Schuster, 1964; Currie, 1976; Gary, 1985, p. 2), but that obviously was not the case on range displayed in photographs in this section.

Johnson (1956) conducted stocking rate trials on the Manitou Experimental Forest and documented changes in plant species composition on ponderosa pine-bunchgrass ranges under different grazing intensities. He found that mountain muhly was the most abundant species on all study pastures, especially under light grazing intensity, and that Arizona fescue was co-dominant with mountain muhly only under moderate grazing. These findings were consistent with observations (and photographs) of the current author. All photographs shown herein were taken on lightly and ungrazed ranges for purposes of showing maximum plant size/development along with best possible expression of vegetation development. Furthermore, climax vegetation (or the closest "best guess" to it) was selected and presented as the benchmark range plant community for examples of the ponderosa pine-bunchgrass range type. It followed then that mountain muhly would be the dominant (most prominent and abundant) grass of range vegetation that was presented in this section.

Perhaps also telling in this context was the fact that mountain muhly but not Arizona fescue was among the 200 species of North American range plants that qualified for the Society for Range Management International Range Plant Contest (Stubbendieck et al., 1992).

The Range Plant Handbook (Forest Service, 1940, G81) put it succinctly: "Because of it great abundance this grass is the most important species in the higher ponderosa pine types in many areas of the Southwest and Colorado". Previous research, agency experience, and current field observations by this author indicating mountain muhly as the dominant of the ponderosa pine-bunchgrass range type was further substantiated by the designation of Pinus ponderosa- Muhlembergia montanaWoodland by the Colorado Natural Heritage Program as listed at end of this caption.

The density and crown cover of ponderosa pine in this range vegetation was of such relative sparcity that it was clearly more woodland or even savanna than forest. Such physiogonomy was evident from these and almost all other photographs in this portion of the chapter on Southern and Central Rocky Mountain forests. The only interlocking of pine crowns occurred in small grove-like groups of, usually, less than ten adult trees. Pine regeneration was also quite limited though certainly adequate to maintain current low densities of uneven-aged pine populations. Dispersion pattern and regeneration of ponderosa pine was shown more specifically in slides below.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

48. Scattered but "sexually active"- Widespread dispersal pattern of trees in a ponderosa pine-mountain muhly woodland did not preclude pine reproduction. A ponderosa pine sapling (left midground, first slide; left foreground, second slide) was featured at edge of a small park (natural "clearing") in the deep interior of this ponderosa pine-bunchgrass woodland. Mountain muhly formed an almost exclusive--though eratic or sporadic(whichever would be more correct)--cespitose coverage of the soil surface ground other than for local single-species assemblages of other grasses (bearded or slender wheatgrass, the closest thing to an associate species on this range) or small stands of mixed species (purple reedgrass, pine dropseed, Nelson's needlegrass).

Recruitment rate of ponderosa pine on this woodland range was conspicuously low, but clearly adequate to maintain the low stand density of this sole coniferous species. Stand density is "a quantiative measure of stocking expressed either absolutely in terms of number of trees, basal area, or vloume per unit area or relative to some standard condition" (Helms, 1998). Clearly, absolute stand density was low (even for ponderosa pine) on this montane woodland. In contrast to stand density, stocking (in a silvicultural context) refers to "an indication of growing space occupancy relative to a predetermined stand" including relative density, basal area, and/or percent occupancy (Helms, 1998). Stocking on this low-tree density woodland was (or ws nearing) 100% of the planned management goal.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

49. Pine grove (sort of)- Exterior of a local aggregation of ponderosa pine showing structure of a mostly eve-aged stand of adult trees. (There were a few mid-size trees behind the foremost three adult pines.) Similar groups of such age/size class trees variously scattered along with isolated individual pines (mostly adults) and "parks" (gladelike patches of bunchgrasses) formed the ponderosa pine woodland range featured in this section. In some places on this range, vegetational structure was more of a savanna.

The closeness of ponderosa pine in the stand shown here was the greatest density of adult trees this photographer could find on this woodland range. Tree crowns barely touched under this closeness or concentration of adult trees.

The apparently most important feature of tree dispersion on this woodland was that distribution of ponderosa pines varied from aggregated to uniform to absence. Dispersion is the pattern of spatial distrbiution of organisms (trees in this instance), the physical arrangement or grouping of individual organisms in the space of their einivonment (ie. pattern of tree occurrence on the land surface of this woodland range). Dispersion ia sometimes called pattern of distribution or simply distribution. This inconsitent pattern of ponderosa pine dispersion was over or across a more consistent herbaceous layer that had relatively uniform structure and a relatively consistent domination by mountain muhly although a few other grass species (bearded or slender wheatgrass, pine dropseed, Indian ricegrass, purple reedgrass) were dominant in local (small, restricted) spots.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

50. Interior structure of a sparse stand (of trees that is)- Structure and species composition of a woodland to savanna form of a montane ponderosa pine-bunchgrass range in the interior of the Southern Rocky Mountains.Pines were of such wide spacing and relatively sparse cover (low tree density and canopy coverage) that the range vegetation could accurately be described as savanna, at least in some parts of the plant community. The absolute stand density (defined above) of this range vegetation was so low that the herbaceous understorey was well-developed across the range (throughout the plant community) with large, robust plants of the various grass species.

The herbaceous layer(s) was featured immediately below. The larger-scale perspectives presented here included a number of grass species including mountain muhly, the dominant, New Mexico sleepygrass, slender or bearded wheatgrass, purple reedgrass, Nelsons needlegrass, Indian ricegrass, blue grama, and,undoubtedly (though the author could not identify any), Arizona fescue. Forbs, all of which were very sparsely scattered, included wholeleaf Indian paintbrush, Rocky Mountain pussytoes, sagewort wormwood or field sagewort,and Lambert's crazyweed.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

51. The herbaceous (feed) layer(s)- Two "photoplots" of the herbaceous understorey of a montane ponderosa pine-bunchgrasswoodland range in the interior of the Southern Rocky Mountains. In the foreground of the first of these two "photoplots" was a consociation of mountain muhly, the overwhelming dominant grass species of this range cover type (at least in this area). This was an other species-excluding understorey stand of this native, cespitose, eragrostoid grass. The grey-colored forb was sagewort wormwood or field sagewort.

The second "photoplot" was of a more botanically diverse local plant community that included Nelson's needlegrass, pine dropseed, blue grama, sun caric sedge, and Rocky Mountain pussytoes in addition to mountain muhly, the overall and local dominant dominant, and field sagewort. Distinctive and local (small spatial-scale) assemplies of various herbaceous species such as this one in the second had developed throughout the understorey of this woodland range.(Another such assemblage was shown in the very next photograph.) In some of the larger parks and old fields New Mexico sleepygrass formed single-species stands that were also small in area (and example was shown farther below).

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

52. Other bunchgrasses- Another "photoquadrant" of the herbaceous layer (s) of a montane ponderosa pine-bunchgrass woodland range in the Southern Rocky Mountains. This local community ("mini-community") of herbaceous plants was dominated by Indian ricegrass, but there was considerable species richness and overall botanical diversity in this "photosample"(in contrast to the mountain muhly consociation presented immediately above). Herbaceous species shown here were overwhelmingly grasses including in addition to Indian ricegrass, slender or bearded wheatgrass, Nelsons needlegrass, purple reedgrass, blue grama, and, perhaps Arizona fescue (although the author could not positively identify any of this latter species on this range at time of photography). Also present was sun caric sdege (the only grasslike plant identified in this local sample of range vegetation) and Rocky Mountain pussytoes.

Local (spatially restricted or small-area) herbaceous communities such as this also developed in larger parks located within the surrounding woodland. Some examples of these park "mini-communities" were presented farther below.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

53. Prominent herbaceous citizens- Two large plants of mountain muhly (first slide) and one mid-size and one small plant of mountain muhly (second slide) along with plants of Indian ricegrass, bearded or slender wheatgrass, Nelson's needlegrass, purple reedgrass, Fremont's geranium, and Rocky mountain pussytoes (largely indistinguishable in backgrounds) in the herbaceous layer of a ponderosa pine-bunchgrass woodland range in the Southern Rocky Mountains. Presence of ponderosa pine was evident by a basal trunk (first photograph) and needles on lower branches (second photograph). A portion of the crown of a common juniper (Juniperus communis) was in lefthand-margin of first slide.

These two "photoplosts" were representative of the mountain muhly-dominated herbaceous layer of this potential natural vegetation that was identified and described as by Colorado Natural Heritage Program as Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance all under Southern Rocky Mountain Ponderosa Pine Woodland. It was explained above why mountain muhly was the sole dominant of the herbaceous zone. If a Daubenmire habitat type was named for this forest range vegetation it would have to be ponderosa pine-mountain muhly.

More photographs and further description of mountain muhly were presented below in conjuction with associated understorey species.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year).

54. Younger pines and a little closer together- A second-growth ponderosa pine-bunchgrass forest with an even-aged population of young pines. This stand was contiguous with the uneven-aged ponderosa pine-bunchgrass range in the eight slide-pnoto caption sets described immediately above. The forest range vegetation seen in these two slides had higher stocking and a greater proportion (higher percentage of relative cover) of jponderosa pine. There was a much higher proportion (relative cover) of Arizona fescue (though mountain muhly was still the most abundant grass) on this forest range with greater tree density than in the forest range vegetation described above that had lower lower tree density (lighter stocking) and greater relative cover (more complete dominance) of mountain muhly.

This stand of ponderosa pine was under a shelterwood regeneration method which is one type of even-aged management. In the shelterwood method most of trees of initial reproduction following the last harvest are cut, usually in a sequence, leaving a stand composed of one age class that cast enough shade to result in a moderate forest environment (Helms, 1998). IThe forest stand seen here was in effect a plantation of ponderosa pine. It was part of a long-term, in-depth study involving analysis of growth and yield of even-aged pines within the pole-size class along with growth and development of the understorey across different stockings (Gary, 1985, ps. 10-13).

Herbaceous species in addition to mountain muhly and Arizona fescue included pine or hairy dropseed, slender or bearded wheatgrass, blue grama, purple reedgrass, and Nelson's needlegrass as more important grasses along with such common forbs as sagewort wormwood or field sagewort, .Rocky Mountain pussytoes, Lambert crazyweed, and one-sided penstemon. In isolated spots little bluestem (Andropogon scoparius) was important. Presence of this tallgrass species along with blue grama, a shortgrass species, and the midgrasses just listed demonstrted the affinity of this forest-bunchgrass range community with mixed prairie on the adjoining Great Plains and Colorado Pateau. In fact the understorey of this forest community was mixed prairie.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year).

55.Denser and more prolific ponderosa pines- A forest form (greater absolute density, closer spacing) of uneven-aged ponderosa pine that developed in proximity to (roughly half a mile from) the woodland and/or savanna form of ponderosa pine-dominated community featured above. This was a different forest site that was much more mesic. Reproduction was much greater in this range plant community than in the wide tree spacing (low density, sparse canopy cover) ponderosa pine-bunchgrass range shown previously. Small saplings and large seedlings of Colorado blue spruce (Picea pungens) were interspersed with those of ponderosa pine. Blue spruce was represented only by large seedlings. Blue spruce served as an indictor species of wetter soil. Another additional species in this ponderosa pine forest was quaking aspen, some of which were visible in the immediately succeeding slide. Quaking aspen and blue spruce are sometimes associated on more mesic sites. A detailed account of that relationship was described elsewhere in this and the quaking aspen chapter..

Species composition and structure of the understorey under ponderosa pine with greater density and more nearly complete canopy cover was quite similar to that of the savanna and woodland forms except for presence of far more forb species and less dominance by mountain muhly in the forest form of this range type. Composite or overall species richness and botanical diversity (including structure as well as species composition) was visibly greater in the forest form of this cover type.Grasses found on this "photofield" included mountain muhly, bearded or slender wheatgrass, Nelson's needlegrass, pine dropseed, Indian ricegrass, plus muttongrass or Fendler's bluegrass (Poa fendleriana) which was not found on the more open canopy forms of ponderosa pine-bunchgrass. Conversely, New Mexico sleepygrass was not in and blue grama was much less abundant in the forest form of ponderosa pine-bunchgrass (and there would be no missing the three to five foot-tall sleepygrass).

Pike National Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37)..Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

56. Where ponderosa pines grew a little thicker (and another tree species chummed in)- Forest (vs. woodland or savanna) form of ponderosa pine-bunchgrass range in the Southern Rocky Mountains with a local and sporadic shrub or lower tree layer of quaking aspen. There were saplings to small-pole size quaking aspen as well as seedlings and/or suckers (clonal shoots) from these. Only seedlings of aspen were present in the first of these two photographs whereas saplings and one seedling of aspen were visible in the second photograph.

The first photograph presented the interior structure and composition of a ponderosa pine-bunchgrass range community and, in the foreground, one of the few local habitats that was not under tree canopy. The second photograph featured the herbaceous layer under a dense local stand of adult ponderosa pine. Forest vegetation shown in the second slide was in the distant background of the first photograph.

The dense, bright-green carpet in the first slide was colonies of Fendler's groundsel or Fendler's ragwort in the phenological stage prior to bolting or emergence of sexual shoots (flower stalks). These shoots can extend to foot-tall (or greater) heights. On this range the few early flowering plants had sexual shoots of about six to nine inches height (shown below with other plant species).

Grass species composition in the herbaceous layer(s) of this forest range were more diverse than in the woodland or savanna form that was dominated overwhelmingly by mountain muhly. Under denser ponderosa pine there was greater relative cover and density of such grasses as bearded or slender wheatgrass, pine dropseed, Nelson's needlegras, and Indian ricegrasss than where less crown cover was a factor in dominance by mountain muhly. Sun cric sedge was not found in this denser (more canopy shade) ponderosa pine forest.

Forbs in the ponderosa pine-bunchgrass forest range shown here (as well as in the preceding slide) included Fendler's groundsel or Fendler's ragwort that was shown at a distance in the first photograph, one-sided penstemon, Fremont's geranium, Rocky mountain pussytoes, and sagewort wormwood or field sagewort. Forbs such as wholeleaf Indian paintbrush, Lambert's crazyweed, prairie thistle, and miner's candle that were common on parks and more open-canopy ponderosa pine woodlands were absent from this closed canopy (or close to it) forest.

The herbaceous layer on the forest floor presented in the second slide was dominated by mountain muhly showing that this overall dominant herbaceous species could "hold the ground" even under fairly dense shade. Rocky Mountain pussytoes and Fremont's geranium were conspicuous forbs in this scene.

Pike National Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37)..Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

57. Where pines grew a lot sparser- A large park situated within the general range plant community of ponderosa pine-bunchgrass woodland or forest. In the parlance of Landscape Ecology such parks are patches within the matrix of ponderosa pine-dominate forest. The application of the term park was explained in detail below at conclusion of the treatment of the ponderosa pine-bunchgrass type in the Front Range of the Southern Rocky Mountains.

The most abundant species on this park was mountain muhly, but other important (even locally dominant) grass species included pine or hairy dropseed, New Mexico sleepygrass, slender or bearded wheatgrass, blue grama, purple reedgrass, and Arizona fescue. Sun caric sedge was also common (and locally dominant). Forbs were rare (usually non-existant), but the two most common ones in this early summer vegetational society were miner's candle (Cryptantha virgata) and prairie thistle (Cirsium canescens).

Pike National Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). There appeared to be no vegetational unit for such parks within the general ponderosa pine-bunchgrass range ecosystem.

58. Park sward- Closeup of the range vegetation on the large park introduced immediately above. The dominant was mountain muhly (cespitose plants in right foreground and left midground) which was also the dominant herbaceous species of this general range plant community which was ponderosa pine-mountain muhly woodland (Southern Rocky Mountain Ponderosa Pine Woodland- Colorado Natural Heritage Program). The other major grass species on this large park were pine or hairy dropseed (bunchgrass in left foreground), blue grama (several sexual shoots in immediate foreground), and New Mexico sleepygrass. Arizona fescue was present, but clearly not a major species. Sun caric sedge was the most abundant non-grass species on this park though none was visible in this photograph.

Pike National Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). There appeared to be no vegetational unit for such parks within the general ponderosa pine-bunchgrass range ecosystem.

59. Mounded mountain muhly- Examples of mountain muhly (Muhlenbergia montana) on a park situated within a ponderosa pine-bunchgrass woodland range in the Southern Rocky Mountains. Previous photographs in this section presented examples of mountain muhly in "photoplots" of the herbaceous layer(s) immediately under ponderosa pine. Plants of mountain muhly seen in these three slides were on an unshaded park were quite a bit larger and more robust than those beneath pines (ie. in forests or woodland versus on parks).

In addition, these three photographs were taken at closer camera range (focal distance) including one from a more top-down view (first photograph). The tufted habit of this strictly cespitose (all shoots are tillers) species was evident in both these and the two slides shown above. Also conspicuous in all five of these photographs was the cascade-like or drappingpattern of broken-down leaves (and a few culms) that formed a thatch at base of plants. This is partly the result of leaf sheaths that extend beyong shorter subtending internodes and curled basal leaves (Barkworth et al., 2003, ps. 184-185).

Sources cited above unequivocally showed that mountain muhly was either the most abundant and important herbaceous species (Forest Service, 1940, p. G81; Johnson, 1956) or a co-dominant perennial grass (Schuster, 1964; Currie, 1976; Gary, 1985) in ponderosa pine-dominated forests or what the Colorado Natural Heritage Program (undated) titled the Pinus ponderosa- Muhlembergia montanaWoodland unit of Pinus ponderosa Woodland Alliance.

In addition to these references pertaining to features of this species, including forge value, readers were further referred to Shaw (2008, ps. 157-159) and Stubbendieck et al. (1992, ps. 124-125). The latter source was devoted to the 200 range plant species included on the Society for Range Manageament International Range Plant Contest. Inclusion of mountain muhly among the "SRM 200" bespeaks importance of this species.

Mountain muhly is widely distributed in North and Central America with a species range extending from Wyoming to Guatemala (Barkworth et al., 2003, p. 184). Palatability of mountain muhly is high to fair for most species when ahoots are at less mature vegetative stages, but herbage acceptability to grazers is much less at advanced maturity, especially with dead or dormant shoots (Foresst Service, 1940, p. G81). Like most grass species that grow to larger plant size, smature hoots of mountain muhly become rank and the herbage of this species "stemy" in later phenology.

Pike National Forest, Teller Counter, Colorado. Early July; mid-growth, pre-boot stage of phenology.

60. Mounted clusters- In this two-slide set and the set immediately following, panicles of mountain muhly were shown at progressively closer views. The first slide presented upper portions of shoots of a single plant of mountain muhly with emphasis on two sexual shoots complete with just-emerged panicles. These two panicles with their flag leaves were displayed at closer camera range in the second slide. The flag leaf of grass is the leaf immediately subtending (below) the flower cluster (ie. the flower cluster, a panicle in Muhlenbergia species, and the culm bearing it emerges from the sheath of the uppermost leaf which is known as the flag leaf).

Pine straw (shed pine needles) from ponderosa pine littered the ground on which this mountain muhly plant grew. This was most clear in the first slide which also included a ponderosa pine cone visible in left rear margin. The most conspicuous neighbor plant was Fremont's geranium.

Pike National Forest, Teller Counter, Colorado. Early July; advancing stages of flowering.

61. Mounted closer- Closer-up details of the flower cluster of mountain muhly. The first of these two photographs presented a single panicle emerging (almost fully emerged) from the boot with the large flag leaf in front of the panicle. The second photograph was an attempt to show details of panicle branching and spikelets in mountain muhly.

The spikelets of the Muhlembergia species are relatively small; those of M. montana are some of the larger ones, especially when open during anthsis.

Pike National Forest, Teller Counter, Colorado. Early July; advancing stages of flowering

62. Another ergrastoid grass- Pine, beardless, or hairy dropseed (Blepharoneuron tricholepis) in the herbaceous layer of a ponderosa pine-bunchgrass woodland range in the Front Range of the Rocky Mountains. This cespitose, C4 pathway grass is locally abundant and regarded as one of the most nutritious and palatable grass species on forest ranges throughout the Southern Rocky Mountains south through the Sierra Madre. Feed value declines fairly dramatically with maturity and, even more so, dormancy (Forest Service, 1940, G 24).

Importance of this eragrostoid grass is attested to by its inclusion in the Society for Range Management list of 200 North American range plant species for the International Range Plant Identification Contest (Stubbendieck et al., 1992, ps. 114-115). .Pine dropseed was described in the Range Plant Handbook (Forest Service, 1940, G 24) has being about equal in forage value to mountain muhly with which it grows in association on open forest ranges.

The involute basal leaves, especially those of the previous growing season, are a readily identifying characteristic of this species (of course, curled or twisted leaves are a feature of several grass species). This characteristic was most evident in the third slide in this set. Blepharoneuron is represented in North America by two species, the other of which is an annual found in Mexico. Blepharoneuron is closely related to Sporobolus and, in fact, B. tricholepis was formerely included in that genus as S. tricholepis. There is one other Blepharoneuron species in North America It is an annual native to Mexico.

Pike National Forest. Early July- peak bloom.

63. Slender samples- Examples of slender or bearded wheatgrass (Agropyron subsecundum= A. trachycaulum). Here we go again. For decades agrostologists distinguished between bearded wheatgrass (A. subsecundum) and slender wheatgrass (A. trachycaulum) as, for example, in Hitchcock and Chase (1950, ps. 231, 238) where the dichotomy was between spikelets awnless or awn-tipped only (A. trachycaulum) and spikelets awned (A. subsecundum). The plant species list for the Manitou Experimental Forest (Gray, 1985) included both of these species. At time this caption was written, slender or bearded wheatgrass was designated as Elymus trachycaulus while A. subsecundum was not recognized as being in Colorado (Weber, 1990, p. 268; Shaw, 2008, ps. 582-583).

It has long-been accepted that "beauty is in the eye of the beholder" ( apparently from the Greek before the time of Christ to final form by Margaret Wolfe Hungerford). It now seems that species and othr taxa are in the eye of the taxonomist. World-renowned agrostologists from Hitchcock and Chase (1951) to Barkworth et al. (2007) can look at slender and/or bearded wheatgrass and see not only different species but different genera (Agropyron versus Elymus). "Whatever" (Archie Bunker).

Likewise, whatever scientific (or common) name is used, slender or bearded wheatgrass is regarded as excellent forage for livestock and wildlife, plus it was interpreted in North American Range Plants (Stubbendieck et al, 1992, ps.204-205) as having either awnless or awned lemmas. The latest edition of this reference/identification guide used Elymus genus, but given that the word "range" was replaced in the title of this work this author refused to cite it.

Bearded or slender wheatgrass is strictly a bunchgrass (ie. all shoots are tillers; it lacks both rhizomes and stolons).

Pike National Forest, Teller Counter, Colorado. Early July; early bloom stage.

65. Mature specimens-Several sexual shoots with mature (grain-ripe) spikes (first slide) and detail of a mature spike (second slide) of bearded or slender wheatgrass on a ponderosa pine-bunchgrass woodland range. All of these spikes had prominently awned lemmas so that they keyed to Agropyron subsecundum in the old taxonomy (Hitchcock and Chase, 1950) or to A. trachycaulum after these two Agropyron species were merged or to Elymus trachycaulus after cladistics turned taxonomy on its head (Skinner et al., 1999).

Pike National Forest, Teller Counter, Colorado. Early July; soft-dough stage of grain.

66. Needling Nelson- Nelson's needlegrass or subalpine needlegrass (Stipa nelsonii= Stipa columbiana var. nelsonii) or, due to confusion, Columbia needlegrass (S. columbiana= S. columbiana var. nelsonii) in the understorey of a ponderosa pine-bunchgrass woodland range in Southern Rocky Mountains.This species was one of the grasses that--along with slender or bearded wheatgrass, pine dropseed, Indian riecgrass, purple reedgrass, blue grama, Arizona fescue, muttongrass or Fendler's bluegrass,and bottlebrush squirreltail--was associated with mountain muhly, the dominant grass species. Some of these bunchgrasses, mostly Indian ricegrass and mountain muhly, were in the background of the first slide.Also present was sun caric sedge and various forbs. These forbs were featured below after the grass species.

Both of the individual plants of Nelson's needlegrass in these two slides (one plant per slide) exemplified the cespitose (tufted) or bunchgrass habit. Both of these plants had this year's pre-boot shoots along with last year's shoots and their shed-out infloresences.

Nelson's or subalpine needlegrass has been regarded as one of the more important Stipa species in the 11 public lands states-portion of the Western Range. It was the Stipa species singled out for inclusion in the Range Plant Handbook (Forest Service, 1940, G115).

There has been considerable confusion regarding nomenclature of this taxon. Under the traditional (Linnean) taxonomy S. nelsonii (rather than S. columbiana) was finally decided upon (Weber, 1990, p. 285). Previous literature such as the plant species list for Manitou Experimental Forest (Gary, 1985) used S. columbiana, but the older (historically useful) authorities, as for example (Coulter and Nelson, 1909, p. 49), had distinguished between (based on length of lemma awn for example) and described both S. columbiana and S. nelsonii. Hitchcock and Chase (1950, p. 458) recognized S. columbiana var. nelsonii based on larger plant size and longer awns. The Range Plant Handbook (Forest Service, 1940, G115) also used the designation, S. columbiana var. nelsonii. To add further confusion S. nelsonii and/or S. columbiana were confused with S. dorei. Skinner et al. (1999, ps. 127-128) distinguished between S. nelsonii subsp. nelsonii and S. nelsonii subsp. dorei.

Finally (as of this writing) this species (or species complex) was transferred to another (different) genus based on cladestic criteria (Barkworth et al., 2007; Shaw, 2008, p. 520) which the current author did not see fit to record and dignify. Cladist authors retained the two subspecies epithets nelsonii and dorei.

Pike National Forest, Teller Counter, Colorado. Early July

67. Last year's glory- Sexual shoots with panicles of Nelson's or subalpine needlegrass from the previous year (growing season) that had overwintered and retained their spikelets.Obviously it had been a dry, mild winter. Drier conditions were still in effect at time of this and above photographs so that even in early July current year's shoots were still in boot or even pre-boot stages.

Pike National Forest, Teller Counter, Colorado. Early July; grain-ripe stage (why they were not shed after this time was unknown.

68. Robust, but sleepy- Specimens of sleeepygrass or, sometimes in full, New Mexico sleepygrass (Stipa robusta= S. vaseyi) at edge of a small park within a ponderosa pine-bunchgrass woodland range. Sleepygrass is an infamous toxic species that is lengendary in history and folklore as well as in Range Science, Toxicology, and other sciences. The common name is based on the fact that this species is toxic to livestock, especially horses, with symptoms being extreme stupor lasting up to a few days after a poisoning incident.

The main toxic compound appears to be ergine, a lyserbic acid amide (LSA) which is an alkaloid produced by a fungal endophyte in the species in genus Acremonium (= Neotyphodium), possibly A. chisosum (Kaiser et al., 1996; Glenn et al., 1996). LSA is similar to the synthetic hallucinogen LSD. LSA occurs in greatest natural concentrations in sleepygrass (Halpern, 2004, p. 135). Petroski et al. (1992) reported four other alkaloids in addition to LSA in S. robusta. Acremonium is the same genus of fungal endophte living in tall fescue (Festuca arundinacea) and respondible for an array of fescue-toxicity symptoms. White (1987) found that the sleepygrass endophyte appeared to be sexually sterile so that its survival depended exclusively on asexual reproduction. Sleepygrass toxicity is obviously an interesting, if not intriguing, phenomenon.

The definitive authorities for stock-poisoning by sleepygrass are Kingsbury (1964, p. 493-494)--also noted in this work's predecessor text (Muenscher, 1961, p. 40)--and the current "bible" of North American poisonous plants by Burrows and Tyrl (2001, ps. 945-947). The latter concluded that while some poisoning still takes place, even rarely resulting in death, "[s]leepygrass is now more of a toxicologic curosity than a significant disease problem".

One of the earliest reports of the narcotic-like effect of sleepygrass on livestock and, again, especially in horses was in the Flora of New Mexico (Wooton and Stanley, 1915, p. 67), a work of great scholarship that itself became a legend.

Given the rich lore of sleepygrass in the territory and, later, state of New Mexico this species is more commonly associated with this political unit, hence New Mexico sleepygrass. There is even a Sleepygrass Campground on the Lincoln National Forest a short distance from Cloudcroft, New Mexico where numerous samples of sleepygrass have been collected for chemical analysis (eg. Petroski et al., 1992).

For the ponderosa pine-bunchgrass woodland type in the Southern Rocky Mountains Gary (1985) summarized studies on natural revegetation (secondary succession) on old fields in the Manitou Experimental Forest which concluded that sleepygrass was a seral species. Wooton and Stanley (1915, p.67) noted that sleepygrass "grows vigerously and spreads rapidly, especially when other plants are killed by overstocking". This generally fits the textbook example of an increaser species. Similarily, Wolfe (1979) reported that sleepygrass eincreased for a period with overgrazing of more preferred range plants, but that it was at a competitive disadvantage to several other grass species. Interestingly, Jones et al. (2000) found that cattle exhibited preference for endophyte-free sleepygrass and speculated that the alledged avoidance of sleephygrass by livestock was a response to presence of endophytes more than to minor diffrences in nutritional value of forage.

Pike National Forest, Teller Counter, Colorado. Early July; early vegetative stage(s).

69. Tall sleeper in the woods-Three-slide sequence of panicle through spikelets of sleepygrass that was growing on a small mountain park in ponderosa pine-bunchgrass woodland range. These panicles (as well as sexual shoots in the preceding two-slide set) were last year's material that had weathered a mild and dry autumn through early summer period with little, if any decomposition. Weather variables such as wind, rain, and hail had not been strong or severe enough to disperse ripe florets and caryopses that had been produced the preceding year.

At this point in time sleepygrass was still in roughly mid-stages of vegetative growth while other cool-season, C3 grass species like Indian ricegrass and purple reedgrass (Calamagrostis purpurescens) and even some plants of warm-season, C4 grass species including mountain muhly were in phenological stages ranging from anthesis through soft-dough states. Jones et al. (2000) explained that the rank-growing sleepygrass had "very late maturity" and that, although a cool-season species, was still in strictly vegetative stages when other cool-season grasses had already senesced.

Pike National Forest, Teller Counter, Colorado. Early July; dead panicles (grain-ripe stage) from the previous year.

70. Sun caric sedge (Carex heliophila= C. inops subsp. heliophila= C. pensylvanica subsp. helipophila= C. exiebenia)- Portions of three plants of sun sedge (first photograph) and details of sexual shoots of sun sedge (second slide) on a small park within an overall ponderosa pine woodland to forest-bunchgrass range type in the Southern Rocky Mountains. These plants had already entered dormancy in a drier than typical spring-summer growing season.

Sun caric sedge is a local dominant and one of the most abundant/important Carex species, especially on drier microsites, in the ponderosa pine-bunchgrass cover type (Gary, 1985).

Pike National Forest, Teller County, Colorado. Early July; dormant phenological stage.

71. Thistle in the park- Prairie or Platte thistle (Cirsium canescens) growing on a small park situated within an overall ponderosa pine woodland bunchgrass range in the Southern Rocky Mountains. Unlike many introduced and subsequently naturalized Cirsium species on North American ranges C. canescens is a native forb and not one inclinded to become weedy (at least not a major invasive species).

Prairie thistle is frequently found on parks in the ponderosa pine forest zone as well as on mixed prairie (Weber, 1990, p. 80), but it is not a major species.

Pike National Forest, Teller County, Colorado. Early July; at peak phenological stage.

72. Plants and a panicle- Portions of five plants of Indian ricegrass (Oryzopsis hymenoides) growing in a consociation of this species that comprised a semidesert grassland in northern San Luis Valley.Second photograph was a part of a panicle of one of these plants showing the tremendously heavy grain crop produced in an extremely wet growing season. These plants and those shown in the immedately preceding photograph of the above pair of slides were giant specimens of their species growing on pristine (Excellent range conditin class) semidesert grassland that was made up almost exclusively of this one species.

Indian ricegrass growing in uderstories of ponderosa pine-bunchgrass forest or woodland typically do not grow to such immense size. The author made these photographs do double-duty to show habit of this remarkable species which is the State Grass of both Utah and Nevada.

Bureau of Land Management, Foothills Allotment. Alamosa County, Colorado. Mid-June, hard-dough phenological stage.

73. Grainy details- Mid-season culms and leaves and just-emerged spikelets of Indian ricegrass growing in the herbaceous layer of a ponderosa pine-bunchgrass woodland range in the Southern Rocky Mountains. Neighboring species included mountain muhly, purple reedgrass, pine dropseed, slender or bearded wheatgrass, and Nelson's or subalpine needlegrass among the Gramineae, Rocky Mountain pussytoes and sagewort wormwood or field sagewort as the major forbs, and Arkansas rose as the sole shrub species.

Pike National Forest, Teller Counter, Colorado. Early July; anthesis.

74. Making grain for the squaws- Spikelets of Indian ricegrass with exerted anthers in the herbaceous layer of a ponderosa pine-bunchgrass woodland range in the Southern Rocky Mountains. These spikelets were in late anthesis as readily determined by anther exertion on lower flowers. Grasses are determinate bloomers meaning that flowering (ascertained by anthesis) commences in the upper and outer (versis lower and inner) florets and progresses downward and inward.

.Typically, panicle branches are paired with, in turn, two spikelets per branch Shaw, 2008, p. 517). Spikelets of Indian ricegrass have only one floret with disarticulation above the glumes Lemma awns are shed early-on in the grain ripening process.

The common name, Indian ricegrass, was derived from the aboriginal practice of squaws and Indian maidens gathering its grains (florets and caryopses) and then winnowing and finally ground into a meal or coarse flour which used for bread (Pavlik, 2008, ps. 3-7). This practice was particularily widespread southwestern desert tribes such as the Paiute, Mojave, and Navajo.

Recently workers at Montana State University developed receipes for flour made from the gluten-free and relatively nutritious grain of Indian ricegrass. They then convinced some traditional grain (eg. wheat)-producers growers to start growing Inidan ricegrass for specialty or niche markets. Now the Amazing Grains Grower Cooperative specializes in production of Indian ricegrass grain for "health food" markets (reporter J.Stromnes, Missoulian, 16 November, 2003).

Pike National Forest, Teller Counter, Colorado. Early July; advanced anthesis.

75. Purple pinegrass under ponderosa pine- It might not improve one's enuniciation to repeat those words ten times in rapid fire, but it will enhance his understanding of the botanically rich herbaceous understorey of ponderosa pine-bunchgrass woodland range to learn about purple pinegrass or purple reedgrass (Calamagrostis purpurescens) on this forest range type.

The first of these three photographs showed a small or local colony of purple reedgrass whereas the second slide included two plants (one behind the other) and the third presented only one plant of purple pinegrass. All of these plants were ion the same ponderosa pine-bunchgrass forest (woodland form) range in the Southern Rocky Mountains.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year; prebloom stage but with dead and westhered sexual shoots from the preceding year).

76. Fremont's floral footprint- Fremont's or pinewoods geranium (Geranium caespitosum subsp. caespitosum vr. fremontii= G. fremontii= G. parryi) in the herbaceous zone of a ponderosa pine-mountain muhly forest range in the Southern Rocky Mountains. This important forage species has been the subject of much debate in regards it proper nomenclature and, even, identity.

This is one of the most widespread range forbs in Rocky Mountain Region. The closely species, G. richardsonii, made the Society for Range Management list of 200 range plant species for the International Range Plant Identification Contest (Stubbendieck et al., 1992). G. richardsonii is the major Geranium species on the west (Pacific) side of the Continental Divide and into the inland Northwest. G. fremontii is the east-side equivalent although both of these species grow--often together--on the Atlantic side of the Big Divide. In point of fact, these and even a third species, G. atropurpureum, frequently hybridize (Weber, 1990, p. 203).

Pike National Forest, Teller, County, Colorado. Early July; peak bloom stage of phenology.

77. Fendler's groundsel (Senecio fendleri)- The herbaceous composite was one of the more common--hence, important--forbs in the uncerstorey of a ponderosa pine-mountain muhly forest/woodland range. This forb was more common on the denser, closed-canopy forest form more than on the woodland form of the general ponderosa pine cover type.

Pike National Forest, Teller, County, Colorado. Early July; full-bloom stage of phenology.

78. Pussy footing under the pines- Rocky Mountain pussytoes (Antennaria parvifolia) on floor of a ponderosa pine-mountain muhly forest range. This composite grew on the lowest leve of the herbaceous zone (or lowest herbaceous layer). It compositious neighbor was Fendler's groundsel.

Pike National Forest, Teller, County, Colorado. Early July; full-bloom stage of phenology.

79. On the floor and off to the side- One-sided penstemon (Penstemon unilaterialis= P. virgatus subsp. asa-grayi) of the needle-littered floor of a ponderosa pine-bunchgrass woodland range in the Southern Rocky Mountains. This plant was not a robust specimen that is more typical of its species where there is more light and moister soils.

Pike National Forest, Teller County, Colorado. Early July; full-bloom phenological stage.

80. One-sided details- Closer-in views of the inflorescence of one-sided penstemon. This specimen was growing on a subalpine mountain meadow that was subirrigated by lateral flow of water from Big Thompson River. One-sided penstemon is adapted to a diverse array of habitats ranging from the more shaded and drier floor of ponderosa pine-bunchgrass woodland ranges to parks within ponderosa pine forests to subalpine mountain meadows.

Membership in the snap-dragon family (Scrophulariaceae) was only all-too-apparent in this view of flowers of one-sided penstemon.

Larimer County, Colorado. Early July; peak full-bloom stage of phenology.

81. Lodgepole lupine- An individual--and a comparatively large--plant of Lupinus parvaflorus growing in the herbaceous zone of a ponderosa pine-bunchgrass woodland range. The local habitat of this particular plant was a pine needle-covered floor where pines were growing at greater density (more like that of a forest). This denser growth of trees resulted in canopy cover much greater than that of the more common woodland form of this range plant community in which crowns of individual pines seldom come into contact with each other.

This lupine apparently is not poisonous to livestock.

Pike National Forest, Teller, County, Colorado. Early July; full-bloom stage of phenology.

82. Standing milkvetch (Astragalus adsurgeus var. robustior)- This was another papilionaceous legume that was locally abundant in the herbaceous vegetation of a ponderosa pine-mountain muhly woodland range shown above.

Although many Astragalus species are infamous stock-poisoning range plants A. adsurgeus var. robustior has long-been regarded as nontoxic. It does contain swainsonine at concentrations high enough to put animals consuming it at risk. Nonconsumption of standing milkvetch may explain the fact that it apparently is not poisonous to livestock (Burrows and Tyrl, 2001, ps.518-519).

Manitou Experimental Forest, Teller Counter, Colorado. Early July.

83. One out of the textbooks- Lambert's crazyweed or Lambert's locoweed (Oxytropis lambertii) is one of the textbook examples of a poisonous range plant. Incidence (frequency) of poisoning, colorfulness of plant, and early publicity (not necesaarily the good kind) have marked Lambert's loco as one of the classic stock-poisoning plants.

This was the first species described and pictured (twice and in color) in Stock-Poisoning Plants of the Range (Marsh, 1918).It was in the colored frontispeace as a principal stock-poisoning plant in Range and Pasture Management (Sampson, 1923), the first textbook in Range Management. Durrell and Newsom (1936) described poisoning by this species (known at that time as Aragallus lambertii) on Colorado ranges. Of course O. lambertii was in Kingsbury (1964, ps. 307-311) and, with expanded coverage, in Burrows and Tyrl (2001, ps. 514, 594-599 passim), the current "bible" of toxic North American plants.

Manitou Experimental Forest, Teller Counter, Colorado. Early July.

84. Very pretty, and even more poisonous- Individual papilionaceous flowers of Lambert's crazyweed. Attractive plant, and historically one of the deadliest ones to livestock. Lambert's locoweed causes classic locoism in cattle, sheep, and horses. lThe poisonous principle in Lambert's locoweed that induces locoism is the same as in Astragalus species: swainsonine and similar compounds. Unlike Astragalus species, however, those of Oxytropis are not selenium accumulators. On the other hand, some Oxytropis species (eg. O. sericea) cause right-heart failure commonly called "big brisket" ,"brisket disease", and "high elevation sickness".

Details of swainsonine, a indolizine alkaloid, and its toxicity were provided in Burrows and Tyrl (2001, ps. 518-522). More details can be found in wikipedia, the free encyclopedia.

Manitou Experimental Forest, Teller Counter, Colorado. Early July.

85. Brushed beneath the pines- Wholeleaf Indian paintbrush (Castilleja integra) in the herbaceous layer of a ponderosa pine-bunchgrass woodland range in Southern Rocky Mountains. This species was one of the more common forbs on drier, more sunlite microsites.

Manitou Experimental Forest, Teller Counter, Colorado. Early July.

86. Arkansawer in the Rockies- Arkansas rose (Rosa arkansana) in lower shrub layer of a ponderosa pine-bunchgrass woodland range in Southern Rocky Mountains. Rosa species are somewhat of a complicated group (sort of a hair-splitting situation), but the bristles on the shoot, nine leaflets or more per leaf, and the rounded non-tapering hips give this species away. (Besides R. arkansana was the only rose on the species list of the Manitou Experimental Forest [Gary, 1985].)

This plant was growing in the company of mountain muhly, pine dropseed, Indian ricegrass, bearded or slender wheatgrass, Nelson's needlegrass, purple reedgrass, Fremont's geranium, and Rocky Mountain pussytoes.

Manitou Experimental Forest, Teller Counter, Colorado. Early July.

87. Looking rosy- Details of inflorescence and leaves of Arkansas rose in understorey of a ponderosa pine-bunchgrass forest (a woodland form) range in the Southern Rocky Mountains.

Manitou Experimental Forest, Teller Counter, Colorado. Early July.

88. A hip plant in the woods- Several hips on a plant of Arkansas rose in a ponderosa pine-bunchgrass woodland range. Hip is the false-fruit of Rosa species where false-fruit refers to the structure or organ formed from merger of the separate true fruits of (Smith, 1977, see Glossary therein). The false fruit or Rosa species, the rose hip, is actually the tough, thick hypanthium formed from the floral cup or floral tube.

Manitou Experimental Forest, Teller Counter, Colorado. Early July.

89. Landscape - Landscape-scale view in interior of Southern Rocky Mountains showing ponderosa pine-bunchgrass forest and woodland in the distant background, a park within ponderosa pine woodland in foreground, and a wide zone of riparian vegetation (mostly a willow carr) along a mountain stream in the midground. The ponderosa pine-bunchgrass range community was covered immediately above and below whereas the willow carr was covered in less detail below at end of the ponderosa pine-bunchgrass woodland range type.

From perspective of Landscape Ecology the ponderosa pine-bunchgrass (mountain muhly being the dominant bunchgrass species) woodland was the matrix of this vegetational mosaic while parks within the woodland were patches and the stream, Trout Creek, was a corridor. Matrix is the largest vegetation unit of organization within a landscape (as the term is applied in Landscape Ecology); it is the encompassing areal or regional vegetation. Patch refers to plant-determined communities (animals can be included) inside of (embedded within) the matrix (eg. parks within area-wide woodland or forest). Corridor is the organizational unit, including its vegetation, of a landscape that passes through matrix and, sometimes, patches. Corridor can connect patches or even different matrices. Streams (ranging in size from brooks to rivers) are probably the most common and important (critical) kinds of corridors. Other corridors include passes through mountains, game trails, and such human-constructed corridors as railroads and highways, irrigation cannals, and power line corridors.

The park in the foreground wass a consociation of mountain muuhly with very other grass species present. These incidental species included blue grama, Nelson's needlegrass, and sun caric sedge. Forbs were, for all practical purposes, absent.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37)..Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland (Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

90. Park in the pines- It was explained at various points throughout this chapter that park as applied throughout parts of the Rocky Mountains simply means any naturally occurring opening of herbaceous vegetation within a surrounding (typically a larger) forest or woodland. A park can be quite small such as just a portion of an acre to an large or even giant-sized area of thousands of acres such as the famed North Park, Middle Park, and South Park in the Colorado portion of the Southern Rocky Mountains. Park is a term long-established in the vocabulary of mountain folk who use park in a generic sense to mean native vegetation devoid (or largely so) of trees or shrubs (ie. a natural "clearing"). Park in reference to land supporting only or, at least mostly, herbaceous plant life (ie. grassland, marsh, mountain meadow, etc.) goes back to the earliest days of entry into these parts by white man--explorers and mountain men.

The comparatively small park shown in these two photographs and the one presented immediately above were within the surrounding (and area-wide) natural vegetation of ponderosa pine-bunchgrass (mountain muhly, the dominant) woodland. The park displayed here was dominated by New Mexico sleepygrass so named for its well-known and early documented (Wooton and Stanley, 1915, p. 67) stupfying narcotic effect on horses. Studies on the Manitou Experimental Forest reported that sleepygrass was more abundant on abandoned croplands (ie. "go-back land") where it was subclimax to the mountain muhly-Arizona fescue climax (summarized by Gary, 1985, ps. 2-3). To the south in New Mexico, Wooton and Stanley(1915, p. 67) described sleepygrass (Stipa vaseyi shown therein) as: "A very abundant grass in meadows at middle elevations". Wooton and Stanley :(1915, p. 67) explained further that locally raised livestock did not eat sleepygras sso that S. vaseyi grew robustly and spread quickly, in particular when other range plants were killed out by overgrazing. Grazing on the park shown here was limited to wildlife, especially mule deer. Perhaps range animals on this park avoided sleepygrass. This author certainly found no evidence of grazing, but degree of use was almost nil for every other plant species.

Other grasses included the almost ever-present mountain muhly, pine dropseed, blue grama, purple reedgrass, Nelson's needlegrass, and bearded or slender wheatgrass. Sun caric sedge was also present, but (and in contrast to an adjjacent park only a few hundred yards distant) there were almost no forbs preent except for field sagewort.

Conifers around margin of this park were, of course, ponderosa pine.

Pike National Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). Apparently no vegetational unit for such parks. General range vegetation was as followed. FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37)..Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland (Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

91. Park residents- Two "photoquadrants" of natural range vegetation on a park within surrounding ponderosa pine-mountain muhly woodland. The first photograph featured pine dropseed, the foremost cespitose plant, with slender or bearded wheatgrass (behind and to right of the pine dropseed), sun caric sedge, and blue grama also present. The second "photoplot" held a diverse array of species including mountain muhly (the most abundant but not greatly so), pine dropseed, New Mexico sleepygrass, blue grama, sun caric sedge, and bearded or slender wheatgrass.

This range vegetation was on still yet another park (a different one from the two described in the two immediately preceding captions). This park had been grazed by cattle in the not-too-distant past as proven by numerous heaps of cattle dung such as the prominent one in the center foreground of the second slide (dung was also present in background of the first slide, but can be seen clearly only with projection of the slide).

Pike National Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). Apparently no vegetational unit for such parks. General range vegetation was as followed. FRES No. 21 (Ponderosa Pine Forest and Woodland Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine). No SRM but would be a southern variant of SRM 110 (Ponderosa Pine-Grassland). Pinus ponderosa Association 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37)..Pinus ponderosa-Muhlembergia montanaWoodland within Pinus ponderosa Woodland Alliance (Southern Rocky Mountain Ponderosa Pine Woodland (Colorado Natural Heritage Program). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

92. Down Trout Creek-Mountain stream and broad valley in the Southern Rocky Mountains surrounded by ponderosa pine-mountain muhly woodland and forest. In context of the landscape concept Trout Creek was a wet corridor through a matrix of ponderosa pine-bunchgrass forest with numerous relatively small parks that functioned as patches in the forest matrix.

The riparian vegetation of Trout Creek was primarily a carr of coyote willow (Salix exigua) with occasional plants and colonies of thinleaf alder (Alnus incana subsp. tenuifolia), water birch (Betula occidentalis= B. fontinalis), and shrubby cinquefoil (Potentilla fruticosa). Smooth brome (Bromus inermis) was about the only herbaceous species growing around these shrubs as visible in these two photographs. Smooth brome is a naturalized species which-- while an extremely agronomic species for tame pasture, various range types, and high-yielding hay--has widely displaced many native species at numerous locations.

There are a myriad of Salix species throughout the Rocky Mountain Region (and adjoining environments). Weber (1990) listed over 20 native species or distinctive subspecies of Salix for Colorado's Eastern Slope. Positive identification of most of these species usually requires both male and female catkins as well as fruits. Local workers may be familar enouth with the willows (and throughout the year) to be able to identify and distinguish species by association of vegetative features with those of fruit and flower. The rest of us have to rely on short cuts such as the handy species list for the Manitou Experiment Forest (Gary, 1985) which in this instance quickly limited willows shown here to S. exigua.

Streams (regardless of size) are of much more value for water yield, watershed protection, water for wildlife, habitat for fish, and sites for riparian vegetation which also have unique values.

Manitou Experimental Forest, Teller Counter, Colorado. Early July (early estival aspect in a dry cool year). No FRES or K- Nos. SRM 422 (Riparian). SAF 235 (Cotton-Willow). Willow Series 231.71 in Rocky Mountain Alpine and Subalpine Swamp and Riparian Scrub biotic community 231.7 of Brown et al. (1998). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

93. A common willow- Several plants of coyote willow (Salix exigua) in the riparian zone of Trout Creek in the Southern Rocky Mountains. All plants in these two photographs were females. The Salix species are conspicuously clonal plants with numerous shoots arising from the rootcrown and resulting in a cespitose habit.The second of these two photographs illustrted that feature.

Willows are often heavily browsed by various kinds of range animals, including cattle. In fact, where range cattle hang in on streams willows are usually overbrowsed, sometimes to point of extirpation. Overbrowsing and elimination of willows can occur just as readily from elk though they are usually less apt to "camp out" on willow carrs. The willows seen here were subject to browsing by mule deer, but not elk or cattle. There was no evidence of "highlining", elimination of almost all browse within reach of range animals resulting in a prominent lower layer on browse plants that is largely devoid of leaves or, even, shoots entirely.

The herbaceous layer of the limited length of riparian zone shown here was dominated (almost exclusively on some microhabitats) by naturalized smooth brome.

Manitou Experimental Forest, Teller Counter, Colorado. Early July; mature fruit just prior to shedding.

94. Particulars of coyote willow- Three progressively closer views of the leader of a female coyote willow. Most of the branch that bore leaves was shown in the first slide whereas specifics of leaves and female catkin were given in the second and third photograph. Willows, like cottonwoods, are dioecious species.

Manitou Experimental Forest, Teller Counter, Colorado. Early July; mature fruit just prior to shedding.

In the southeastern portion of the Colorado Plateau (more specifically, the San Francisco Plateau Plateau of the Grand Canyon section within the Colorado Plateau physiographic province) and at the edge of the Mogollon Rim (Mogollon Plateau) above Verde Valley there is the largest conterminous ponderosa pine or western yellow pine (Pinus ponderosa) forest in North America. This ponderosa pine forest includes numerous range sites and habitat types as well as subtypes, variants, or forms of the generic ponderosa pine forest range type designated as forest cover type, SAF 237 (Interior Ponderosa Pine) by the society of American Foresters (Eyre, 1980).

In the area centered around the neat or "nifty" human community of Flagstaff, Arizona the author found examples of both 1) the ponderosa pine-Arizona fescue range type and 2) ponderosa pine-mountain muhly range type. Gambel's oak (Quercus gambelii), as--both or either--shrub and tree and numerous valuable browse plants were also major components or members of this assemblage of forest range vegetation.

Examples of this forested range plant community were presented in the following section.

As was the case for several other units of the ponderosa pine forest cover type treated in this chapter bearing the designations of Southern and Central Rocky Mountains, the Arizona ponderosa pine type (or subtype, variant of SAF 237) is not in the Rocky Mountain System, either Southern Rocky Mountain or Middle Rocky Mountain physiographic provinces (Fenneman, 1931, ps. 92-182; Thornbury, 1965, ps. 322-383). The floristic affinity of the Arizona ponderosa pine forest--like that of the Black Hills (Great Plains physiographic province) and those of the Basin and Range physiographic province--to ponderosa pine-dominated forests in the Rcoky Mountain system was such that it was deemed appropriate to include the "outliers" of the Rocky Mountain chain herein under the designation (for lack of a better one) of "regionally related" forest range types.

95. Ponderosa pine-Arizona fescue (Festuca arizonica) range- An example of the physiogonomy and simple structure of the Arizona form of ponderosa pine forest with consociation of Arizona fescue as understorey. This was the three-needle fascicle Rocky Mountain ponderosa pine (P. ponderosa var. scopulorum) and not the five-needle fascicle Arizona ponderosa pine (P. ponderosa var. arizonica). Vegetation was of climax composition and structure, and proved that forest and range resources can be used wisely to produce renewable crops of forage, wildlife (and wildlife habitat), wood, and water.

Coconino National Forest, Coconino County, Arizona. Vernal aspect (June), but severe to exceptional drought so that phenological development of fescue was delayed. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

96. Structure and species composition of Arizona form of parklike ponderosa pine-Arizona fescue forest- This photograph showed the local (= micro-) topography or landform of a ponderosa pine forest cover type in the Colorado Plateau of central Arizona. There was adequate regeneration of pine to maintain the open, woodland-like forest stand. The understorey consisted of a single herbaceous layer composed almost exclusively of Arizona fescue (a consociation). At other locations (microsites, perhaps) the exclusive herbaceous understorey was a consociation of Stipa neomexicana (see below). In these single-species herbaceous layers there were scattered plants or, less commonly, local populations of blue grama. These did not seem adequate to warrent interpretation of blue grama as an associate species. Interestingly, mountain muhly (Muhlenbergia montana), which is often an associate or even co-dominant species of this range type, was not found in this vegetation. Cheatgrass (Bromus tectorum) grew in trace amounts, but this was restricted to road berms.

Coconino National Forest, Coconino County, Arizona. Vernal aspect, June (exceptional drought conditions). FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Fores). SAF 237 (Interior Ponderosa Pine), an Arizona variant. No SRM designation for this cover type. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

97. Two "photo-plots" of Arizona form of Rocky Mountain ponderosa pine-Arizona fescue forest range- These two slides revealed a more diverse vegetation than that shown in preceding ones. Vegetation in these "photo-plots" (slides) included a few widely scattered shrubs. In the first slide a shrub element (if it could be called that) was represented by rubber rabbitbrush (Chrysothamnus nauseosus) in the far-right foreground. Immediately in front of the rabbitbrush was a local population of Stipa neomexicana. As shown below, this grass species also formed a single-species herbaceous layer in parts of this ponderosa pine forest. The rest of the understorey was composed of cespitose individuals of Arizona fescue (both green and brown-- still dormant-- tufts).

In the second slide, shrubs (but clearly not a shrub layer) were represented by a species of gooseberry (Ribes sp.) seen at left foreground. The woody plant behind and to the right of the gooseberry was a yound ponderosa pine. Students should observe (in both "photo-plots") that ponderosa pine consisted of several age classes, including small trees, indicating regeneration of this species.

Vegetation shown in these photographs was the natural potential of climax stage. Many of the larger pines had the form of old-growth specimens, but even if a few logs had been "snaked" out and the range grazed by livestock these were benchmark examples of virgin vegetation (and this vegetation was shown after four dry years that rated designation of exceptional drought).

Coconino National Forest, Coconino County, Arizona. June, vernal aspect (severe to exceptional drought). FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. No SRM designation. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

98. Arizona form of Rocky Mountain ponderosa pine-Arizona fescue forest range vegetation- In contrast to the parklike or woodland-like physiogonomy of the ponderosa pine-dominated community shown above (and below), some areas of this forest had more of a vegetational mosaic or patchwork. In this "photo-plot" a natural opening consisted of a "pure" stand of Arizona fescue while an adjoining patch was a stand of young ponderosa pine so dense that it's shade prevented growth of any species beneath it resulting in pine straw-covered ground and no understorey.

The igneous parent material of the soil was portrayed by the conspicuous boulders in foreground.

Coconino National Forest, Coconino County, Arizona. Vernal (and drought) aspect, June. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. No SRM recognition for this cover type. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

Arizona ponderosa pine ranges have several species of grasses and shrubs which may either form biologically diverse understories of one to several layer(s) or, alternatively, "species sparse" understories consisting of a single-species stand or, at most, a few species. The single-species understorey examples of Arizona fescue and New Mexico feathergrass presented here were forms of parklike ponderosa pine forest. Other ponderosa pine forests have more biological diversity as both species-richness and structural complexity (more layers in the vegetation).

Coconino National Forest, Coconino County, Arizona. Estival aspect (but under conditions of exceptional drought), June. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF (Interior Ponderosa Pine), another Arizona variant. No SRM designation. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

Coconino National Forest, Coconino County, Arizona. Estival aspect, June

104. Specimens of New Mexico feathergrass (Stipa neomexicana) in ponderosa pine forest understorey- These robust plants were at seed-ripe or seed-shatter stage in a Rocky Mountain ponderosa pine parklike forest of the Arizona form. Exceptional drought conditions, but there had been enough soil moisture from winter precipitation for these cool-season natives to complete their annual growth cycle "with flying colors".

Coconino National Forest, Coconino County, Arizona. Beginning of dormant season for these denizens, June.

106. Grains of New Mexico feathergrass- Four ripe caryopses (each still enclosed within a palea and lemma, the latter of which has a sharp-tipped callus at base of a characteristically twisted awn) of S. neomexicana. These grains showed the potential for mechanical injury to grazing animals.

Coconino National Forest, Coconino County, Arizona. June.

107. Blue grama (Boutelous gracilis)- Blue grama was an associate of the dominant Arizona fescue and New Mexico feathergrass in the understorey of the Arizona form of ponderosa pine forest. Blue grama is one of the most widespred and important range plants in North America. In fact, if blue grama is second to anything in total acreage and forage production it is little bluestem. Blue grama retains it's nutritive value better in the dormant stage than most other grass species. It's culms and leaves sometimes remain in semi-dormancy which allows the plant to "green-up" and begin photosynthesizing more quickly than those species that have to initiate all new growth from the rootcrown. This is also a major reason why blue grama remains more nutritious during the dormant season. Blue grama is an eragrostoid grass (Eragrostoideae subfamily) rather than festucoid (Festucoideae subfamily) like Arizona fescue and New Mexico feathergrass.

Coconino National Forest, Coconino County, Arizona. Delayed estival aspect due to exceptional drought, June.

108. Slender wheatgrass (Agropyron trachycaulum)- This species is another festucoid grass that is sometimes a local dominant but it did not assume the overall dominance of Arizona fescue or New Mexico needlegrass (= feathergrass) in the understorey of the Arizona form of ponderosa pine forest.

Actually most of the grass cover in this "photoplot" (the lower, tan-colored foliage) was cheatgrass (Bromus tectorum), a naturalized Eurasian annual

Coconino National Forest, Coconino County, Arizona. Peak standing crop; grain in soft dough stage (June).

109. Red threeawn (Aristida longiseta)- Red threeawn is generally an invader on most range types and sites except for desert ranges where it may even be interpreted as a climax grass (eg. in the Sonoran Desert). In the understorey of ponderosa pine forest ranges red threeawn is a "black sheep" of the grass family compared to climax and productive forage species like Arizona fescue, New Mexico feathergrass, mountain muhly, and blue grama. This invader species is, however, of value as an indicator plant (indicative of disturbances including overgrazing) the high population of which serves as a warning to the range manager. The attractive specimens seen here were growing on-- and doing "their part" to stabilize-- a road cut during an exceptional drought (worst since Arizona began keeping records). They were valuable as both an indicator species and soil protector plus provider of relatively unpalatable forage.

Coconino National Forest, Coconino County, Arizona. June.

Vegetation of the ponderosa pine forests of southwestern North America as well as adjoining parts of the juniper-pinon woodland and mixed prairie grasslands commonly have shrub species, some of which provide substantial amounts of palatable browse and thereby serve as indicator species for ecological status of the range and degree of use. Many range plant communities of these cover types have a very limited shrub element and, frequently, no shrub layer (as in the Arizona examples of pristine Rocky Mountain ponderosa pine). Some shrubs are usually present however even if as minor members of the range vegetation. Browse produced by these woody species (thus the species themselves) is of more importance than suggested by "minority" status.

110. Cliffrose (Cowaina mexicana var. stansburiana)- Cliffrose, Mexican cliffrose, Stansbury cliffrose, or quninie-bush is one of the most valuable browse plants in the Colorado Plateau and mountains of the Basin and Range provinces. However it is rather limited within it's distributional range and it's brittle branches are easily broken which can result in low vigor or death of plants even under conditions of otherwise proper degree of use. Cliffrose is rated as good to excellent for browsing wildlife like deer but only fair to poor for cattle. Nonetheless, cliffrose is easily "browsed out" on livestock ranges where animal distribution is improper or inadequate. It was one of the major browse species drastically reduced by overstocking of range animals-- both livestock and wildlife-- during the early part of the Twentieth Century. In fact, the classic (and still controversial) study of grazing capacity and wildlife populations was that of the Kaibab Plateau range and deer herd of northern Arizona in the general area where these photographs of cliffrose were taken. (Students were referred to the abbreviated but readily available discussion by Humphrey [1962, ps. 138-142 passim].)

For general discussion of cliffrose the following timeless standards were recommended: Dayton (1931, ps 49-50), McMinn !1939, p. 204-205), Forest Service (1940, B68), and Sampson and Jespersen (1963, ps.85-87). Good summaries can be gotten from the various editions of North American Range Plants by Stubbendieck, Hatch, and others (1881, 1986, 1992, 1997).

The individuals of cliffrose shown here were mature-- rather old, actually-- specimens in the San Francisco Mountains in the Grand Canyon section of the Colorado Plateau. They were both blooming and bearing fruit. Coconino National Forest, Coconino County, Arizona.

111. Young plant of Mexican cliffrose- This young specimen was growing beside the larger plants shown in the preceding slides. It was not too young to have sexual reproduction. Coconino National Forest, Coconino County, Arizona. June.

112. Fruit-laden leaders of Stansbury cliffrose- Even after four years of the worst drought (rating of exceptional) in history of Arizona weather records the cliffrose plants presented here were flowering while bearing their newest fruit crop. Coconino National Forest, Coconino County, Arizona. June (and into a four-year exceptional drought awaiting summer rains).

113. Inflorescences and fruits of Mexican cliffrose (Cowaina mexicana var. stansburiana)- This species is in the Rosoideae (rose subfamily) of the Rosaceae the members of which have the classic rose hypanthium.The cliffrose fruit is an achene, a dry indehiscent fruit whose pericarp and testa (seed coat) are separate. Other genera of this subfamily that are major browse plants include Fallugia (eg. Apache plume), Cercocarpus (eg. the various mountain mahoganies), Rosa (the many wild roses), Rubus (blackberry, dewberry, raspberry), Purshia (antelope bitterbrush), and Adenostoma (chamise and red shanks). These various plants were discussed variously thoroughout this publication

Coconino National Forest, Coconino County, Arizona. June (into the fourth year of a drought rated as exceptional when this photograph was taken).

Note: Some authorities place this species in the Pursia genus as Pursia mexicana var. stansburiana. Mexican cliffrose does hybridize with the bitterbrush, P. tridentata var.glandulosa (= P. glandulosa).

114. Patch of Apache plume (Fallugia pradoxa)- Even under conditions of an extended (four years), exceptional drought (the worst; highest drought ranking) the native browse plant was "setting the woods afire" with it's colorful fruit. Yes, paradoxically it is the plumose fruits not inflorescences that appear to "bloom" in this valuable (for browse, erosion control, aesthetics, ecological indicator) range plant. Apache plume is generally ranked as fair (more-or-less) in browse rating, but in contrast to restricted species like cliffrose Apache plume is widely distributed and often locally abundant making it more valuable than indicated by it's browse rating.

Standard references again include Dayton (1931, ps. 50-51), McMinn (1939, p. 204), Forest Service (1940, B77), and Stubbendieck and Hatch with others (1981, 1986, 1992, 1997).

Coconino National Forest, Coconino County, Arizona. June.

115. Leaders of Apache plume laden with fruit- Close-in shot of the browse shown in the preceding photograph. The fruit is an achene. Coconino National Forest, Coconino County, Arizona. June.

116. Inflorescences of Apache plume- Stamen and hypanthia of flowers were visible even from this distance. Lincoln County, New Mexico. June.

117. Scrub live oak (Quercus turbinella)- Leaves and fruit (acorn) of one of the dominant scrub oaks (species of shrub oak) in the Arizona (= interior) chaparral and southern ponderosa pine forests. Twigs, leaves, and mast (acorns) of this shrub are valuable for browse (and concentrates in instance of acorns) for numerous species of range animals. For example both the Rio Grande and Merriam's subspecies of wild turkey (Meleagris gallopavo intermedia and M. gallopavo merriami, respectively) are found within the range of scrub live oak. Acorn mast is readily eaten by upland game birds, deer, cattle, and species of the pig group such javelina or collarded peccary (Tayassu tajacu= Dicotyles tajacu) as well as destructive feral hogs (Sus scrofa domestica).

118. Outside, looking in- Physiononomy and structure of a ponderosa pine-Gambel's oak-bunchgrass forest range on the San Francisco Plateau within the Colorado Plateau physiographic province. This specific forest range was an example of the climax vegetation for this forest site (actually several forest sites were included) or variant of the interior ponterosa pine forest cover type (the Arizona or southwestern ponderosa pine form or variant).

Ponderosa pine was the the inland or interior variety Pinus ponderosa var. scopulorum. Gambel's oak was extremely variable as to morphology ranging in habit from huge (as in "humongous") trees down to low-growing shrubs. In fact, the same tree (same genetic individual, genotype) existed as both forms there being numerous clonal clumps arising from larger established "trees" (shoots). No other shrub species were present in this view. On upper, drier slopes there were plants of the suffrutescent species, Bigelow's sage or sagebrush (Artemisia biglovii). This was shown below. Large tree-sized Gambel's oak were visible to immediate right of the tall ponderosa pine.

The herbaceous layer consisted entirely of various species of grasses, most of them eragrostoid or festucoid (of subfamilies, Eragrostoideae and Festocoideae, respectively). There was one to a few individuals of forbs including the naturalized,Eurasian, perennial weed, yellow or common toad-flax (Linaria vulgaris), and the native, perennial composite, Arizona muleears (Wyethia arizonica). In essence: no forbs, no shrubs besides scrub forms of Gambel's oak.

In the view seen here--a lower, deeper soil, more mesic part of the range--the dominant grass was blue wildrye (Elymus glaucus) while slender wheatgrass (Agropyron trachycaulum= A. subsecundum) was the associate grass species. Other major grasses included wolftail (Lycurus phleoides), mountain muhly (Muhlenbergia montana), blue grama (Bouteloua gracilis), and, locally, the introduced (Eurasian) species known as intermediate wheatgrass (Agropyron intermedium).

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

119. Moving inside- Structure and composition of a ponderosa pine-Gambel's oak-bunchgrass forest range at climax state on the San Francisco Plateau. The first slide presented a large ponderosa pine in foreground with (in the background) a large tree of Gambel's oak having a three-pronged (three big vertical limbs) pitchfork-shaped crown with offshoots (clonal shoots) arising from near base of the massive trunk.

The second slide presented a different view with the large ponderoda pine in the first phtograph now in left midground and the large pitchfork-crowned gambel's oak with clonal offshoots in right midground. Other such oak offshoots were at left margin in the second slide. Rather thickly spaced (dense) stands of younger ponderosa pines were in the background of both slides. The same split, fallen log was in both of these photographs.

The dominant herbaceous species was blue wildrye while the associate herb was slender wheatgrass. There were some plants of the introduced intermediate wheatgrass as well as wolftail, mountain muhly, blue grama, and Arizona threeawn. Arizona threeawn was uncommon on this lower, less rocky, and less sloping part of the range than on steeper, rockier, and shallower habitats (see below).

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

120. View from inside- Interior of a climax ponderosa pine-Gambel's oak-bunchgrass forest range in the San Ferancisco Plateau. The forest community shown in these two rather restricted views had very few young pines (though there was probably enough regeneration to maintain current tree density) or Gambel's oak such that this was primarily a two-layered forest. Stands of smaller, younger and more densely populated ponderosa pines were visible in the distant background of these two visual perspectives.

The large, adult pines had reached their peak growth and were just entering the senescent stage of their life cycles. Logs littering the ground included both 1) fallen limbs from crowns of these adult ponderosa pines and 2) trunks of smaller pines that had died for reasons unknown to the author. A fire in the distant past of recent history had burnt over portions of this forest range (shown below), but there were no burn marks on the trunks of any of the beautiful trunks presented in these two "photoplots".

Within the area of these two photographs the dominant herbaceous species was slender wheatgrass while the associate species was blue wildrye. Mountain muhly, wolftail, blue grama, Arizona threeawn, and the agronomic species, intrmediate wheatgrass were present with smaller cover and lower density. There were no forbs except the Eurasian weed, yellow or common toad-flax and the native composite, Arizona muleears.

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

121. Co-dominants at different stages- Young adult ponderosa pine and a clonal clump of Gambel's oak in a ponderosa pine-Gambel's oak-bunchgrass forest range in the San Ferancisco Plateau. The shoots of Gambel's oak appeared to be too immature to bear acorns (still at pre-sexual maturity stage of their life cycle).

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

122. Pals, woody and herbaceous- "Nested photoplots" of slender wheatgrass to the fore of a shoot of Gambel's oak in the shrub and herbaceous layers of a ponderosa pine-Gambel's oak-bunchgrass range in the San Francisco Plateau. It was not determined if this shoot of Gambel's oak was a clonal offshoot (ramet) or a large seedling (a genet or unique genotype).

Coconino National Forest, Coconino County, Arizona. Late July.

123. It can do it either way- A lone seedling of Gambel's oak (and a seedling at considerable distance from the nearest clone of this species) provided an example that Gambel's oak reproduces both sexually (as here) and asexually (as shown immediately above and throughout this section).

This baby plant was on the floor of the ponderosa pine-Gambel's oak-bunchgrass forest range featured here. "Pine straw" (accumulated needles of ponderosa pine) and a small plant of some lupine (Lupinus sp.) along with an unidentified grass completed this snapshot of life on this forest range.

Coconino National Forest, Coconino County, Arizona. Late July.

124. Some of the critical characters- Series of three "nested" (or "telescoped") "photoplots" of the plant community of a ponderosa pine-Gambel's oak-bunchgrass forest range in the San Ferancisco Plateau. This local area of the range that was covered in this section had burned sometime in the distant past. The flame-burnished basal trunk of the large pine attested to that fact as did charred remains of limbs and logs that had been on the ground at time of fire.

Gambel's oak was represented by three shoots in right foreground of the first slide. The brown (dead) leafes in upper crowns of these three shoots was stress from prolonged drought that was at Palmer Index category of Extreme to Extraordinary Drought at time of photographs. There were also Gambel's oak shoots (shorter than the three just mentioned) in the left distant background of all three photographs. The two conifer saplings (foremost one blocked view of the second in the second and third of these "photoplots") to immediate left of large ponderosa pine were alligator juniper (Juniperus deppeana). Alligator juniper is one of the few Juniperus species that resprouts following fire or wood cutting. It was not known if these two alligtor juniper had been present at time of the most recent fire, but such did not appear to be the case, there being no charred juniper trunks.

The dominant herbaceous species in the forest community presented in these three slides was slender wheatgrass. Associate herbaceous species (second most abundant grass) was mountain muhly. Other important (common in this range plant community) grasses included wolftail, sideoats grama, blue grama, and Arizona threeawn. There were no plants of blue wildrye or intermediate wheatgrass on this upper slope. The only forb "to amount to anything" was the the naturalized, Eurasian, perennial weed known as yellow or common toad-flax (Linaria vulgaris), an aggressive, hard-to-control invader (Whitson et al., 1992, ps. 548-549).

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

125. Major herbaceous characters- Herbaceous layer, a sward of cespitose native grasses, of a ponderosa pine-Gambel's oak-bunchgrass forest range in the San Ferancisco Plateau. These two slides were "nested photoplots" of the herbaceous layer shown in the immediately three slides (flame-burnished basal trunk of the big ponderosa pine and lower shoot of the foremost alligator juniper in the first of these two slides served as '"plant landmark").

The herbaceous assemblage in the first slide was a consociation of mountain muhly, overall associate herbaceous (grass) species. The second slide (the "nested plot") featured a large plant of mountain muhly (right) and several plants of wolftail (left).

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

126.Another part of the forest community- A local, rocky, south-facing low slope within a ponderosa pine-Gambel's oak-bunchgrass forest range in the San Ferancisco Plateau. This xeric microhabitat supported a consociation or local population of sideoats grama with a few scrubby, shrub-scale plants of Gambel's oak. This was another part of the same forest range that was featured in this section. This mini-sized south-slope habitat with its sideoats grama-Gambel's oak assembly joined (was immediately adjacent to) the portion of this forest range community featured in the two immediately preceding slide-caption sets.

An even rockier, more xeric microenvironment in this ponderosa pine-dominated forest was presented in the next slide ...

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

127. Where its even rockier- A rocky, shallow, steep, southwestern slope was a harsh local evnironment (microsite or microhabitat) within the same forest range that has been being described. This native vegetation was a ponderosa pine-Gambel's oak-bunchgrass forest range in the San Ferancisco Plateau.

The understorey presented in this "photographic dendrogram" consisted of two local grass-dominated communities: 1) sideoats grama (similar to the one featured immediately above which, incidentally, abutted this "photosample" of range vegetation) and 2) Arizona threeawn (Aristida arizonica). The Arizona threeawn-dominated herbaceous community was the subject of the next slide ...

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

128. Grass on the rocks- Two" photoplots" at two locations on a steep, rocky, shallow-soil microhabitat on which local range plant communities were: 1) in first slide, a mixture of Arizona threeawn (Aristida arizonica) and slender wheatgrass and 2) in second slide, an herbaceous assembly of slender wheatgrass, wolftail (with conspicuous contracted panicles), Bigelow's sgebrush (Artemisia bigelovii), Missouri goldernrod (Solidago missouriensis), and the naturalized Eurasian weed, yellow or common toad-flax.

These two local range communities were on a ponderosa pine-Gambel's oak-bunchgrass forest range in the San Ferancisco Plateau. This is the same forest range that has been the subject--the example--of this part of the section devoted to Arizona ponderosa pine forest range.

Coconino National Forest, Coconino County, Arizona. Late July.

129. Parting interior view- Two views deep inside a ponderosa pine-Gambel's oak-bunchgrass forest range in the San Ferancisco Plateau presenting the composition and internal structure of a climax stage of the Arizona variant of this forest range type (SAF 237, Interior Ponderosa Pine). Small plants of Gambel's oak were present as, for example, in left-central midground of the second of these two slides. Dominant grass species was slender wheatgrass. Major grass species included Arizona threeawn, mountain muhly, wolftail, blue grama, sideoats grama, and blue wildrye. There were no forb species in view of these two "photoplots".

Coconino National Forest, Coconino County, Arizona. Late July. FRES No. 21 (Ponderosa Pine Forest Ecosystem). K-18 (Arizona Pine Forest). SAF 237 (Interior Ponderosa Pine), an Arizona variant. There was no SRM rangeland cover type designation for ponderosa pine forest range in any region except Pacific Northwest. SRM 110 (Ponderosa Pine-Grassland) is as close as possible (eg. Festuca idahoensis vs. F. arizonica as understorey dominant)-- and that is not close enough. Pinus ponderosa Association in Yellow Pine Series 122.621 in Yellow Pine Series 122.62, Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37).

Gambel or Gambel's oak (Quercus gambelii) is a widely distributed species of the white oak group (Leucobalanus subgenus) with a species (biological) range extending from the Chihuhuan Desert of Chihuhua north- and eastward to the plains of South Dakota and west to southern Nevada (Elias, 1980 ps. 329-330). Throughout this species range Gambel oak grows primarily in the range communities of the foothills up through various range types of montane forest vegetation. Gambel's oak is a white oak (subgenus, Leucobalanus).

Gambel's oak more commonly occurs as a shrub. In fact, its occurrence in tree form is apparently so uncommon that neither the standard Dendrology text for North American tree species (Harlow et al., 1979) nor the current "bible" of North American silva (Burns and Honkala, 1990) included Gambel oak. As viewers can see below Gambel's oak does grow to tree shape and size; in point of fact, to immense "oak tree size". It is amazing how the leading authorities sometimes "totally miss it".

In the classic manual of trees in North America, Sargent (1933, p.297) described Gambel's oak, which he labeled Quercus utahensis, as "[a] tree, occasionally 30 feet high, with a trunk 4 inches-8 inches in diameter". Kearney and Peeblies (1960, p. 219) and Vines (1960, ps. 161-162) explained that Gambel's oak ranges in size and shape from thicket-forming shrubs as tall as six and a half feet to trees growing fifty feet in height. Kearney and Peeblies (1960, p. 219) listed Q. submollis as a synonym for Q. gambelii. In the classic Flora of New Mexico Wooton and Stanley (1915, ps. 172-174) described Q. utahensis varying from clump-like shrubs to solitary trees, Q. novomexicana ranging from "a shrub" to "a good-sized tree", and Q. submollis ("a low tree or shrub"); but they explained that Q. gambelii would be the single species of white oak in New Mexico if the "...numerous easily recognizble variants of that type..." were ignored.The most descriptive scientific name of this woody species that varies from a sprawling shrub with colony growth form to masive hardwood tree (the plant now called Gambel's oak) was Quercus confusa as used by soem of the early taxonomists.

There has been but limited study into biology and ecology of Gambel's oak (eg. this species was not included in Silva of North America [Burns and Honkala, 1990]). Ryniker et al. (2006) described what they called "structure of Quercus gambelii communities" which included such features as density and basal area of "trees" and age/size classes, but they examined stands of trees (not communities per se; in fact, these were more like populations of Gambel's oak within woody plant communities). Nor did Ryniker et al. (2006) distinguish between genetic individuals or even note the clonal structure/morphology and colony forming characteristic of Gambel's oak

Most of the research involving Gambel's oak involved control of this species in situations where it has become a noxious (ie. brush) species. A study of this aspect, and one that included a thorough reveiw of the Gambel's oak literature up to that point in time, was that of Engle et al. (1983). Muller (1951) seemed to be the best work regarding the clonal or modular nature of Gambel's oak.

General sources for Gambel's oak included Range Plant Handbook (Forest Service, 1940, p. 12) which was best overall from a range plant/range vegetation perspective. Best coverage from a botanical and general view was, in this author's experience, that of (Elias, 1980, ps. 329-330) with the timeless Vines (1960, ps. 161-162) also giving good summary treatment. Lamb (1989, p. 14) briefly mentioned Gambel's oak, but showed distribution by counties in southwesterrn North America. The author of the current publication found it inexplicable tha Q. gambelii was omitted from both the Forest Service's encyclopedia of United States shrub species (Francis, 2004) and that of U.S tree species (Burns and Honkala, 1990). As shown immediately below the silva of the Coconino National Forest (among numerous othernational forests) felt rebuffed and completely rejected.

130. Its a shrub species; its a tree species- A local group of Gambel's oak ranging from large trees (see immediately below) to clumps of multi-stemmed shrubs. It is commonly accepted that Gambel's oak is a highly clonal woody species that primarily reproduces asexually from a system of lignotubers, vast network of woody rootstocks or rhizomes, thereby being capable of covering extensive areas (eg. an entire hillside). Gambel's oak is much like the associated quaking aspen (Populus tremuloides) is this regard. Apparently, this is surprisinglhy little in the scientific literature regarding this feature of Gambel's oak.

It was highly likely that all shoots (from massive oak "trees" to "scrub shoots") were all offshoots or trunks from subterranean lignotubers and/or woody rhizomes so that they were in effect branches off of underground trunks or limbs of the same genetic Gambel's oak (ie. one genotype). This logical conclusion was an "educated hunch" not a genetically determined fact. There were three "damn big" tree-size trunks and over twenty shrub-size trunks (shoots) in this local group of Gambel's oak. Details of two of the tree-size trunks were presented in the immediately succeeding two slides.

This local grove or colony of Gambel's oak was growing in a general, open understorey ponderosa pine forest community; more precisely, a ponderosa pine-Gambel's oak habitat type with an herbaceouslayer of mixed grass species. Several large ponderosa pines were visible in right background of this photograph.

Coconino National Forest, Coconino County, Arizona. Mid-July.

131. One tree or several? Two big trunks of Gambel's oak in a local group or grove introduced in the preceding slide-caption set. Both of these tree trunks or boles were approximately one and a half feet in diameter (DBH). Bark on them was characteristic of mature bark of Gambel's oak.

A thought-provoking exercise is to ask whether these two trunks were two separate (individual) trees (two genetotypes or genetically distinct genetic individuals from two separate acorns) or if there were two mature shoots of the same tree (the same genotype). The latter was the most probable answer (explanation). Also, the shrub-size trunks (of which there were over twenty in this local group) were most likely other offshoots of the same trtee (same genotically unique plant) that had not reached mature of adult size. Probably all of these clonal units were merely limbs or large distinct branches off of the same lignotuber or subterranean network of rootstocks (= woody rhizomes).

Coconino National Forest, Coconino County, Arizona. Mid-July.

132. Shrubs or baby trees? A clump of numerous shoots (first slide) and a single shoot (second slide) of Gmbel's oak in a lower woody layer of a ponderosa pine-Gambel's oak-mixed grass montane forest. Both the clump and the single shoot were growing about fifteen feet of each other. Was the clump and the individual shoot parts of the same plant (same genetic individual) or where these two separate plants (two genotypes)? Clearly the clump of shoots was the same plant with individual shoots being clones or offshoots from the same lignotuber, but was the individual shoot another clonal unit or was it a small sapling of a genetically distinct (separate or second) genotype (genetically unique tree)?

Without genetic analysis, such as DNA testing ("DNA fingerprinting"), there is no way to know. Gambel's oak has both asexual (an example of a Gambel oak seedling was presented below) and sexual reproducution, but asexual or vegetative reproduction predominates.

Coconino National Forest, Coconino County, Arizona. Mid-July.

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133. Studies in trunks and leaves- Bark and leaves on a large (foot and a half DBH) Gambel's oak (first photograph) and on a sapling-size shoot (second photograph) of Gambel's oak in a ponederosa pine-Gambel's oak-mixed grass montane forest. The large trunk with basal leaves was on one of the large "trees" (offshoots) shown in the second slide immediately above; specifically, the trunk on the right of the horizontal phyotograph. The smaller shoot or bole with cluster of leaves shown at closer distance (second photograph) was probably another offshoot of the same tree, the same genetypic individual.

Viewers can readily see from leaves shown in these photographs that Gambel's oak is in the white oak group (subgenus, Leucobalanus).

Coconino National Forest, Coconino County, Arizona. Mid-July.

134. Young juniper amid pines, oaks, and grass- Three-trunk sapling of alligator juniper (Juniperus dippeana) in understorey of ponderosa pine-gambel's oak-mixed grass montane forest (Pinus ponderosa-Quercus gambelii habitat type).

This open, parklike ponderosa pine forest range joined natural range communities of pinyon pine (P. edulis)-juniper woodland and savanna. In addition to alligator juniper (the least common Juniperus speceis) the other junipers were Utah juniper (J. osteosperma), Rocky Mountain juniper (J. scopulorum) , and oneseed juniper (J. monosperma). Examples of these other three Juniperus species on juniper-pinyon pine woodland range that adjoined this forest were shown in the chapter, Juniper-Pinyon Woodland.

Coconino National Forest, Coconino County, Arizona. Mid-July.

135. Slender shoots under sparse pines- Slender wheatgrass (Agropyron trachycalulum= A. subsecundum= A caninum, at least in part) in herbaceous understorey of a ponderosa pine-Gambel's oak-grass montane forest range in the Colorado Plateau of northcentral Arizona. Slender wheatgrass was the overall dominant grass--dominant herbaceous species, for that matter--especially on shallow upslopes. It was a local associate herbaceous species on lower more mesic habitats where blue wildrye (Elymus glaucus) was the dominant herb. As readily ascertained from these photographs (and the first one in the next set) slender wheatgrass is strictly a cespitose (bunchgrass) species.

The two closely resembling species of bearded wheatgrass (A. subsecundum) and slender wheatgrass (A. trachycaulum) were recognized in traditional treatments such as those by Hitchcock and Chase (1950, ps. 231, 238-239) and Kearney and Peebles (1960, ps. 92-94). A widespread older binomial was A. pauciflorum (Forest Service, 1940, p. G5). Gould (1951, ps. 111-112, 120-121) kept these two taxa distinct but as subspecies of Elymus pauciflorus! In Arizona Range Grasses Humphrey (1960, ps6-7), designated slender wheatgrass as Agropyron trachycalulum but in the next edition Ruyle and Young (2003, p. 60-61) changed slender wheatgrass back to Elymus, but as E. trachycaulus! Welsh et al. (1993, p. 820) also used E. trachycaulus. In the latest definitive--definitive based on cladistic criteria--encyclopedia of North American grasses Barkworth et al. (2007, ps. 321-323) interpreted slender wheatgrass as Elymus trachycaulus subsp. trachycaulus (in contrast to E. trachycaulus subsp. subsecundus to which they applied the common name, one-sided wheatgrass). Did you follow all that?

Through all this FUBAR name change--"musical names"--the common name of slender wheatgrass remained. Of course we "lesser lights" are told how superior scientific names are compared to common names. We can all agree that scientific names are superior for the purpose of getting new publications in the taxonomic literature (and resultant academic promotions).

The pertinent point is that slender wheatgrass is a dominant climax grass of montane forest ranges that furnishes feed rated as Good to Excellent in forage value. It provides outstanding soil cover on important forest watersheds. In ponderosa pine forest ranges slender wheatgrass is one of the major forage-producing range plants for early spring green-up. The species (biological) range of slender wheatgrass (in the strict sense not the entire E. trachycaulus complex described above) extends from deep south Mexico to southern portions of the Prairie Provinces eastward to Newfoundland, north to Alaska and west to the Pacific Ocean. The importance of slender wheatgrass is attested to by its being on the Society for Range Manageament Master Plant list of 200 species for the International Range Plant Identification Contest--and under whatever binomial was fashionable at time of that edition, as for example Agropyron trachycaulum in Stubbendieck et al. (1992, ps. 204-205).

Description and discussion in the Range Plant Handbook (Forest Serviece, ps. G5), again as Agroypron pauciflorum (and still the common name of slender wheatgrass), remained one of the best treatments of this species. A fine summary of slender wheatgrass that was much more current (and a favorite of the current author) was Tilley et al. (2011).

On the pine-oak grass montane forest range treated here, slender wheatgrass had already flowered, set and, then, shed grain. Blue wildrye on this same range was in peak to late anthesis by time slender wheatgrass had passed the grain-shatter stage and was entering senescence. The Extrene Drought (Palmer Scale) in which these plants were growing might well have hastened plant maturity of slender wheatgrass.

Coconino National Forest, Coconino County, Arizona. Mid-July; post grain-shatter stage, perhaps entering drought-induced dormancy.

136. Slender bottoms and tops- Specimens of slender wheatgrass on a ponderosa pine-Gambel's oak-grass mountain forest range showing basal shoots, especially leaves (first photograph) and upper sexual shoots with spikes from which grain had already shed (second photograph). The inflorescence of slender wheatgrass is a spike-like or a very open spike (if there is such a thing) that has more recently been described as a spicate raceme (Tilley et al., 2011). It has traditionally been regarded as a spike (Hitchcock and Chase, 1950, p. 238; Jepson, 1953, p. 101; Welsh et al., 1993, p. 820; Skinner, 2010, p. 422). Skinner (2010, p. 422) specified that the rachis is continuous and thus not breaking like the rachis of the traditional Elymus species. That feature was evident in the second of these photographs.

Although cespitose, slender wheatgrass does, sometimes, have short rhizomes (Jepson, 1953, p. 101; Welsh et al., 1993, p. 820). Most leaves of slender wheatgrass plants are at bases of plants (first slide) Slender wheatgrass is best described as a short-lived perennial (Tilley et al., 2011).

Coconino National Forest, Coconino County, Arizona. Mid-July; post grain-shatter stage, perhaps entering drought-induced dormancy.

137. Blue on the forest floor- Two plants of blue wildrye (Elymus glaucus) growing on the forest floor of a ponderosa pine-Gambel's oak-grass range community in the Colorado Plateau of northcentral Arizona. These two plants were growing on shallow, stoney soil at base of a slope in a forest range on which slender wheatgrass was the overall dominant herbaceous species of the understorey. Wolftail, sideoats grama, and Arizona threeawn were other local associates on this upland forest range.

These two specimens were small plants compared to individuals of blue wildrye found on bottomland range sites in the Arizona Transition Zone.

Blue wildrye is a widespread and valuable forage species across the Western Range Region (and, as far as that goes, across much of North America). For all of its importance as a range plant, however, blue wildrye has received relatively little attention. For example, blue wildrye was not included in the University of Arizona publication, Arizona Range Grasses (Humphrey, 1960; Ruyle and and Young, 2003). Gould (1951, ps. 118-119) did provide taxonomic coverage of blue wildrye. Good general references for blue widlrye as a range plant and forage species included the old standby, Range Plant Handbook (Forest Service, 1940, G53), and Dyer et al. (2005).

Coconino National Forest, Coconino County, Arizona. Mid-July; peak to late anthesis.

138. Blue spikes- Examples of spikes (spike type inflorescence) of blue wildrye growing in the herbaceous understorey of a ponderosa pine-Gambel oak-grass montane forest range in the Colorado Plateau of northcentral Arizona. Gould (1951, ps. 118-119) noted that there is usually only one spikelet per node of the rachis in blue wildrye.

Coconino National Forest, Coconino County, Arizona. Mid-July; peak anthesis.phenological stage.

139. Another member of the understorey- Three cespitose plants of mountain muhly (Muhlenbergia montana) in the herbaceous layer of the ponderosa pine-Gambel's oak-grass montane, forest range being treated in this section. Details of mountain muhly were covered earlier (above) in this chapter so such details were not re-presented in this or the following caption.

Plants seen here had been in Extraordinary Drought until a few days prior to these photographs after which water-starved shoots had made considerable growth. Although mountain muhly is typically a climax dominant grass in open ponderosa pine forest ranges (see again above), on this particular range mountain muhly was dominant only in a few isolated microlocations like the one seen here.

Coconino National Forest, Coconino County, Arizona. Mid-July; early stages of shoot growth and development..

140. More muhly- Two "really big" plants with smaller neighboring plants (first slide) and particulars of a third plant (second slide) of mountain muhly in the understorey of a ponderosa pine-Gambel's oak-mixed grass montane forest range in the Colorado Plateau of northern Arizona.

It was remarked in the preceding caption that this growth had been made in an Extraordinary Drought that had broken with light rains only a few days prior to time of these photographs. The dead tissue still on these plants (eg. straw-colored shoots bearing panicles) was last year's growth.

Mountain muhly has a species range extending from deep into Mexico north to British Columbia but mostly limited to the Rocky Mountain Range. The species has been included on the Master Plant List of 200 species for the International Range Plant Identification Contest held by the Society for Range Management (Stubbendieck et al., 1992, ps. 124-125). "Tried and true" treatment of mountain muhly was presented in the Range Plant Handbook (Forest SErvice, 1940, p. G81).

Coconino National Forest, Coconino County, Arizona. Mid-July; early stages of shoot growth and development..

141. Wolfing in the woods- Several cespitose plants of wolftail (Lycurus phleoides) in the herbaceous understorey of a ponderosa pine-Gambel's oak-mixed grass forest range (a Ponderosa Pine-Gambel Oak habitat type). Wolftail was an associate grass species in this range type in which the local dominant herbaceous species was slender wheatgrass (Agropyron subsecundum= A. trachycaulum).

Coconino National Forest, Coconino County, Arizona. Mid-July; shoot maturity, grain-shatter stage.

142. Arizona on the rocks- Four cespitose plants (three in the first of these two slides) provided examples of Arizona threeawn (Aristida arizonica) that was locally abundant on the ponderosa pine-Gambel oak-grass range being described in this section. On this forest range Arizona threeawn grew on and dominated the shallowest, steepest, tmost rock-strewn microsites. Specimens in these photographs were growing under conditions of Extraordinary Drought and still had last year's inflorescences remarkably intact with almost all spikelets completely unshed along with this year's early season shoot growth (from recent rains). Made for neat examples of this species.

Many of the Aristida species are very similar in morphology and taxonomic interpretations of species vary among various "lumpers" and "splitters". A. arizonica is so morphologically unique that agrostologists from Hitchcock and Chase (1950, p. 477-478) to Barkworth et al. (2003, p. 335) have consistently recognized this species. Aristida more than almost any Gramineae genus (with possible exception of the rosettte Panicum species) are subject to interpretation--usually rather arbitrary at that--and have been rearranged about as much as any North American grass species. A. arizonica is very distinctive with its comparatively small size and compact clumped habit growing on its favored habitat of xeric, shallow, rocky slopes (Gould, 1951, ps. 242-243).

None of the Aristida species are preferred by grazing animals, but the threeawns or wiregrasses are utilized as forage, though more on harsh environments such as desert ranges than on favorable range habitats where more palatable plant species are abundant.

Coconino National Forest, Coconino County, Arizona. Mid-July; early shoot growth stage along with last year's unweathered sexual shoots.

143. Ears among the rocks- A plant of Arizona muleears (Wyethia arizonica) in the understorey of a ponderosa pine-Gambel's oak open upland forest. This specimen was growing in the shallow bed of an ephemeral stream (more like a drainage) in this open, parklike forest range.

In Notes on Western Range Forbs Hermann (1966, p. 347) remarked that palatability of this species was "very low". The specimen presented here had no visible evidence of any defoliation.

Coconino National Forest, Coconino County, Arizona. Late July; fruit-ripe stage..

144. Heads above the ears- Two heads or capitula with ripe achenes (first slide) and topdown details of achenes in a head (second slide) of Arizona muleears. These units or organs were on the same plant that was shown in the immediatlely preceding two slides.

Coconino National Forest, Coconino County, Arizona. Late July; fruit-ripe stage..

145. Lightening-struck ponderosa pine- The ponderosa pine cover type is definitely a fire type. Volumes have been written about the role of fire-- both natural and anthropogenic-- in maintenance of these forest and range ecosystems. The historic fire regime was one on high frequency-low intensity such that commonly there was not a build-up of fuel and development of a fire staircase that allowed surface fires to develop into crown fires.

Lightening strikes are the main ignition source for natural forest fires. This mature ponderosa pine standing sentry atop the local high point bore the obvious mark of a direct hit by lightening that could have lead to a forest fire. Whatever fire conditions existed, whether any fire started by such ignition is a low-intensity surface fire or a holocaustic crown fire would depend on fuel. Even if the forest is as dry as the proverbial tenderbox climax ponderosa pine forests with grassy understories like those dominated by Arizona fescue and New Mexico feathergrass are much less likely to become "hell on earth" than those with numerous layers of vegetation leading staircase-like to the forest canopy.

This is the lesson of prescribed burning and proper grazing management on ponderosa pine ranges. Some grazing by livestock and/or big game species reduces buildup of vegetative material that can serve as fuel. Undergrazing of the forest understorey can contribute to destructive crown fires, but so can overgrazing. Prologued overuse that leads to range deterioration leaves the land with less cover as protection against soil erosion. Perhaps even more important in immediate outcome is weakening of the climax grasses which leaves them at a competitive disadvantage against woody plants. This results in increases in shrubs and trees which permits development of the fire staircase as well as leaving the soil surface with less herbaceous growth to break the impact of rain drops or hold back melt water. Understorey management is a key component in proper management-- including wise use-- of fire as a component of ponderosa pine ranges.

Looked at in this perspective, the more-dead-than-alive pine with the "blowed off" crown and the lightening blaze is "a thing of beauty".

Coconino National Forest, Coconino County, Arizona. June.

146. Another form or subtype of the ponderosa pine forest range type is this one in the inland Pacific Northwest where the herbaceous understory layer is dominated by bluebunch wheatgrass and the shrub understory dominants are snowberry and antelope bitterbrush. This is a transition vegetation type or broad ecotone in the Columbia Basin between Palouse Prairie and the ponderosa pine zone of the Blue Mountains and Okanogan Highlands. In the classic Natural Vegetation of Oregon and Washington Franklin and Dryness (1973, p. 172-180) identified both ponderosa pine/ bluebunch wheatgrass and ponderosa pine/ bitterbrush/ bluebunch wheatgrass associations as transitions "between steppe or shrub-steppe and forest". Interior ponderosa pine type. Spokane County, Washington. FRES No. 21 (Ponderosa Pine Ecosystem). K-10 (Western Ponderosa Pine Forest).SAF 237 (Interior Ponderosa Pine). Yellow Pine Series of Brown et al. (1998).

Technical explanation: this example from the Northern Rocky Mountains was inserted here to show 1) the floristic affinity of vegetation of the Northern Rocky Mountains with those of the Central and Southern Rocky Mountains and 2) the continuity and variation within the general ponderosa pine type.

148. Interior Douglas fir forest range type growing on a bottomland or alluvial site along the Snake River— Ponderosa and limber pine (Pinus flexilis) are associated upperstory species. The herb layer is dominated by Poa, Festuca, andStipa species. Transition life zone of C. Hart Merriam. Targhee National Forest, Lincoln County, Wyoming.

FRES No. 21 (Douglas Fir Ecosystem).K-11 (Douglas-fir Forest). SAF 210 (Interior Douglas-fir). Pseudotsuga menziesii-mixed conifer Association in Douglas Fir-White Fir (Mixed Conifer) Series Series 122.61 of Rocky Mountain Montane Conifer Forest biotic community 122.6 of Brown et al. (1998). Middle Rockies- Mid-Elevation Sedimentary Mountains Ecoregion, 17g (Chapman et al., 2003).

149. Mesic form of Rocky Mountain mixed conifer type- In the moist valley of Middle Saint Vrain Creek a mixed conifer forest developed in which Douglas-fir and Colorado blue spruce (Picea pungens) were co-dominant and white fir (Abies concolor) was the overall associate. Lodgepole pine was abundant and a local associate species though more commonly a distant fourth-place conifer to the other coniferous trees. The most common angiosperm tree species were Rocky Mountain willow (Salix monticola) closest to the stream and quaking aspen and Rocky Mountain maple (Acer glabrum) on higher ground farther from the actual riparian zone. Tthinleaf alder (Alnus incana subsp. tenuifolia) and water birch (Betula occidentalis= B. fontinalis) were associate species to Rocky Mountain willow in the range plant community of the riparian zone.

Herbaceous species were mostly limited to the riparian zone and a less shaded margin between riparian and the fully formed mixed conifer forest with nearly complete closed canopy. Herbaceous species included wooly or hairy arnica (Arnica mollis) and common blankeflower (Gaillardia aristata).

Roosevelt National Forest, Boulder County, Colorado. Early July (estival aspect). Elevation: about 8000 to 8500 feet. FRES No. 21 (Douglas Fir Ecosystem).K-11 (Douglas-fir Forest). SAF 210 (Interior Douglas-fir). Pseudotsuga menziesii-Mixed Conifer Association 122.613, Douglas-fir-White Fir (Mixed Conifer) Series 122.61 of Rocky Mountain Montane Conifer Forest biotic community 122.6 (Brown et al., 1998, p. 37). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

150. Conifer mix along a mountain stream - Rocky Mountain mixed conifer community that developed in the floodplain of Middle Saint Verain Creeek. Douglas-fir and blue spruce were co-dominant while white fir was the asociate in most locations with lodgepole pine the associate species at fewer and smaller local environments. Along the streamside margin (riparian zone) of this conifer-dominated forest Rocky Mountain willow was dominant with thinleaf alder and water birch lesser species.Within this Douglas-fir-mixed conifer forest quaking aspen and Rocky Mountain maple were the most important deciduous tree species.

Blue spruce were readily distinguished by their light, bluish or silverish needles. Although blue spruce usually tend to be smaller trees the tall and rather immense conifers in the first photograph were Colorado blue spruce. These remarkable trees were of recordbreaking-like dimensions. Douglas-fir are generally the largest conifers in bottomland habitats in this region, but in this creek bottom forest blue spruce held that distinction. An example of a typical large Dopuglas-fir was shown in the next slide-caption set.

There was almost no understorey in this shade-dense, closed canopy, mesic bottomland forest. There were several herbaceous species in a transition zone between the riparian willow-alder-birch scrub and outer margin of the mixed conifer forest. Most abundant of these herbs were the forbs wooly or hairy arnica and common blankeflower.

Roosevelt National Forest, Boulder County, Colorado. Early July (estival aspect). Elevation: about 8000 to 8500 feet. FRES No. 21 (Douglas Fir Ecosystem).K-11 (Douglas-fir Forest). SAF 210 (Interior Douglas-fir). Pseudotsuga menziesii-Mixed Conifer Association 122.613, Douglas-fir-White Fir (Mixed Conifer) Series 122.61 of Rocky Mountain Montane Conifer Forest biotic community 122.6 (Brown et al., 1998, p. 37). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

151. Moist and mixed- Species composition of a mesic, montane mixed conifer forest that developed in the valley of Middle Saint Vrain Creek in the Suthern Rocky Mountains.Douglas-fir are typically the largest individual trees (height, DBH, crown size) in bottomland coniferous forest in this region, but, as shown immediately above, blue spruce held that distinction in this particular forest. The more typical case of Douglas-fir being the species of the largest trees was reflected in this photograph by the large crown at far left. The other conifers to right of this large Douglas-fir were (left to right): blue spruce, Douglas-fir, white fir, and lodgepole pine.

Also abundant in this conifer-dominated forest were smaller trees of Rocky Mountain maple and quaking aspen. Quaking aspen and blue spruce are often associated in more mesic habitats and forest sites in this general area. That relationship was shown at various points in this chapter as in the section devoted to quaking aspen and above in the ponderosa pine-bunchgrass range type.

There was but a very limited herbaceous component in this closed-canopy bottomland forest. Most herbaceous species were in a transition zone between the riparian willow-alder-birch scrub and outer edge of the dense mixed conifer forest. Some of the more showy forbs in this zone were wooly or hairy arnica and common blankeflower.

Roosevelt National Forest, Boulder County, Colorado. Early July (estival aspect). Elevation: about 8000 to 8500 feet. FRES No. 21 (Douglas Fir Ecosystem).K-11 (Douglas-fir Forest). SAF 210 (Interior Douglas-fir). Pseudotsuga menziesii-Mixed Conifer Association 122.613, Douglas-fir-White Fir (Mixed Conifer) Series 122.61 of Rocky Mountain Montane Conifer Forest biotic community 122.6 (Brown et al., 1998, p. 37). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

152. Slyvan lineup- Two photographs of the major species of a montane, mixed conifer bottomland forest along Middle Saint Vrain Creek in the Southern Rocky Mountains. In the first photograph trees were (left to right): blue spruce, Douglas-fir, and lodgepole pine. The second photograph was a closer-in shot of the Colorado blue spruce and Douglas-fir. All three trees were bearing heavy cone crops.

Roosevelt National Forest, Boulder County, Colorado. Early July. Roughly 8000 to 8500 foot elevation.

153. Big bud on a big tree- New, soon-to-open bud on Douglas-fir. This was not a terminal bud composed of apical meristem arising from the terminus or apex of a branch or leader. Instead it a lateral bud arising from intercalary meristem. This intercalary bud will develop into a new or younger branch off of the older branch rather than growing as a node-internode extension of the older branch as would be the case with a terminal bud.

Roosevelt National Forest, Boulder County, Colorado. Early July. Roughly 8000 to 8500 foot elevation.

154. Slyvan maiden- Immature female cone of Douglas-fir. Gymnosperms (members of Gymnospermae) are seed-bearing, but not flowering plants. Instead of having reproductive structures borne in flowers that mature as fruits as in the angiosperms, the gymnosperms have naked seeds that are produced on unisexual (male only; female only) units known as strobili (strobilus, singular) commonly called a cone (either fleshy or woody). The cone or strobilus consist of a series of sporophylls, which are interpreted spore-bearing, modified leaves, arranged on a common, continuous axis (Smith, 1977, ps. 30, 309). Male cones are smaller and are designated as the microsporangiate strobili. Female cones, the larger of the two sexes, are the megasporangiate strobili. Male and female strobili bear microsporphylls and megasporophylls, respectively. These respective cones are sometimes referred to as staminate strobili and carpellate strobili (Holman and Robbins, 1939, ps. 536-544).

Featured in this photograph of the terminus of a Douglas-fir branch was an immature carpellate or megasporangiate strobili or, in simplier terms a female cone. The author found it a fascinating bit of Botany, and not a bad shot of it either. One of the best explanations--along with outstanding line drawings and clear photographs--of sexual reproduction in the Pinaceae of the Gymnospermae was that of Holman and Robbins (1939, ps. 536-554).

Roosevelt National Forest, Boulder County, Colorado. Early July. Roughly 8000 to 8500 foot elevation.

155. Colorado blue- Leaves and nearing-maturity cones of Colorado blue spruce. There is some variation in colortion of needles of blue spruce, but the glaucous, grey-blue coloration seen here is typical.

Roosevelt National Forest, Boulder County, Colorado. Early July. Roughly 8000 to 8500 foot elevation.

156. Maple sample- Leaves and fruits of Rocky Mountain maple (Acer glabrum). Leaves of this Acer species are clearly "mapleish" and at the same time distinctive from other species being broader and smaller than most of the other Acer species.

The fruit of maples have been interpreted alternatively as either as either a schizocarp or samara.. Smith (1977, p. 165) offered the options, "... as either two samaras, separating at maturity, or a samaroid schizocarp, separating into two-winged mericarps at maturity". Smith defined samara as a dry fruit type that is indehiscent with a single seed and an attached conspicuous wing-like appendage (ie.a winged achene) and schizocarp as a dry indehiscent fruit originating from a syncarpous gynoecium splitting at maturity into one-seeded closed segments called mericarps (Smith, 1977, ps. 66, 307).

Roosevelt National Forest, Boulder County, Colorado. Early July.

157. Range forb in an intermediate zone- Wooly or hairy arnica (Arnica mollis) growing in a transition zone between the riparian willow-alder-birch scrub and outer margin of the mixed conifer forest that was featured immediately. This was mesic habitat, but beyond the actual riparian zone. This composite grew beside the composite featured in the next two slides.

Hermann (1966, ps. 291-292) found that palatability of this composite was variable with ratings ranging from poor or even zero through fair to good. .No such mixed reviews as to its simple beauty.

.Middle Saint Verain Creeek, Roosevelt National Forest, Boulder County, Colorado. Early July.

158. Another forb (and another composite)- Common blankeflower (Gaillardia aristata) on a transition zone between a riparian area that supported a willow-alder-birch shrubland and the leading edge of a mixed conifer bottomland forest dominated by Douglas-fir and Colorado blue spruce and with white fir as the associate species.

Although some Gaillardia species are annuals this showy species is a perennial. It is an extraordinary species for use in landscaping with native plants. Hermann (1966, p. 315) stated that the heads of this range forb were "eaten regularly by cattle and sheep...", but that it was more frequently grazed earlier rather than later in the grazing season.

Middle Saint Verain Creeek, Roosevelt National Forest, Boulder County, Colorado. Early July.

159. Still yet another forb- Whole shoot of sulphur-flowered wild-buckwheat (Eriogonum umbellatum) in first slide and details of the inflorescence of this species in second slide. The plant in first photograph was growing on a higher, drier portion of a subalpine mountain meadow in the Southern Rocky Mountains. Closer view of the flower cluster of this species (second slide) was of a robust plant growing on the local ecotone between a willow-alder-birch srubland and outer edge of Douglas-fir-blue spruce-dominated bottomland forest.

Larimer County, Colorado and Roosevelt National Forest, Boulder County, Colorado, respectively. Early July; peak bloom.

160. Willows within- Rocky Mountain willow (Salix monticola) dominated the riparian zone of Middle Saint Verain Creeek that ran through the mesic, montane mixed conifer forest of Douglas-fir, blue spruce, white fir, lodgepole pine, and Rocky Mountain maple described above.From the perspective of Landscape Ecology this stream and its riparian zone was a corridor within a matrix of the Douglas-fir--blue spruce-dominated floodplain forest.

The zone of Rocky Mountain or park willow was a long, narrow willow carr. Most such carrs (including those with cottonwoods along with willow) are considerably wider than was the case for this thin line of park willow.

The diversity of range vegetation within this area was remarkable. A wide zone of Rocky Mountain lodgepole pine (Pinus contorta var.latifolia) forest developed immediately above the general Douglas-fir forest zone.

The second slide offered a better view of a single plant of Rocky Mountain willow. This plant was a female. Catkins on this plant were presented in the next set of two slides.

Roosevelt National Forest, Boulder County, Colorado. Early July; peak bloom.

162. Furry females- Female catkins of Rocky Mountain or park willow. These structures were on the same plant that was shown in the preceding photograph. The fruit of the Salicaceae is a capsule with comose seeds.

Leaf bases seen here were atypical.

Roosevelt National Forest, Boulder County, Colorado. Early July; peak bloom.

163. General exterior views of a widespread type- Rocky Mountain mixed conifer type in the upper montane zone. This distinctive forest range type is a transitional forest community with botanical elements from the ponderosa pine and Douglas-fir forests, Gambel oak scrubland, and pinyon pine-juniper woodland that are all common range types in the Southern Rocky Mountains. It is a species-rich range plant community with various well-developed and prominent layers. Dick-Peddie (1993, ps. 58, 62-64) described this forest community as the White Fir--Douglas-Fir--Ponderosa Pine Series noting that this type constitutes "the most widespread mixed conifer forests in New Mexico".

In these two "photographic dendrograms" all the taller trees were Douglas-fir except the two leftmost tall (adult to near-adult age/size) trees in the first slide which were white fir (Abies concolor). All of the conifer saplings "on the front row" of both photographs were white fir. The trees in lower-right corner and the extreme right margin of first photograph was quaking aspen as were those in distant midground ar right margin of second slide. There were no ponderosa pine visible in these two slides. Also not visible in these photographs was limber pine (Pinus flexilis) although this species was also well-represented in this forest tract as an associate conifer (see slides below). This was a second-growth forest at an elevation of 9400 feet.

There was regeneration of the three dominant and the associate coniferous species. Most of the herbaceous layer of the understorey was made up of the naturalized, perennial, Eurasian pasture grass, smooth brome (Bromus inermis) as shown in the right foreground in the second of these slides. Other grasses included the natives mountain bromegrass (B. ciliatus, probably as varieties genuinus and/or intonsus) and, almost assuredly here and throughout this area, fringed brome (B. canadensis= B. richardsonii= B. ciliatus var. richardsonii). In fact, many authorities lump these simply as B. ciliatus. Individuals of these (this) species were deep in the interior of this stand and had not progressed to anthesis as had the ubiquitous smooth brome. kentucky bluegrass (another naturalized, perennial, Eurasian species) was common especially around edges and in more open sun-lite spots. There were perhaps other Poa species, but at their pre-bloom stage of development this author could not identify them. There were a few plants of what--at earlier phenological development--appeared to be Idaho fescue (Festuca idahoensis). Of course, there were some plants of cheatgrass, but the perennial bromes, including smooth brome, and Kentucky bluegrass "outweighed" cheatgrass by quite a bit. (Incidentially this author chose not to dignify cladistic classification by showing names based on what he viewed as obscure, arbitrary genetic features.)

The herbaceous layer also included Sey's or prickly rose (Rosa acicularis subsp. sayi). In this layer was the low-growing shrub known as low Oregon grape, holly grape, or low holly grape, (Mahonia repens= Berberis repens).

North La Veta Pass (9400 feet), Costillo County, Colorado. Late June (estival aspect). FRES No. 21 (Ponderosa Pine Ecosystem). Mapped as Kuchler- 17 (Ponderosa Pine-Douglas-fir Forest). Some inconsistency here because Kuchler unit 17 is the only unit that that includes both ponderosa pine and Douglas-fir, but K-17 was listed under Ponderosa Pine Forest Ecosystem not Douglas-Fir forest Ecosystem. SAF 210 (Interior Douglas-Fir). White Fir-Douglas-Fir-Ponderosa Pine Series under Upper Montane Coniferous Forest (Dick-Peddie, 1993, ps. 62-64). Rocky Mountain Montane Conifer Forest 122.6, Douglas Fir-White Fir (Mixed Conifer Series) 122.61, Pseudotsuga menziesii-Mixed Conifer Association, 122.613 in Brown et al. 91993, p. 37). Southern Rocky Mountains- Sedimentary Subalpine Forest Ecoregion, 21b (Chapman et al., 2006).

164. On the outside and going inside of a mixed conifer forest- Second-growth upper montane forest of Douglas-fir, white fir, ponderosa pine, limber pine, quaking juniper and common juniper (Juniperus communis) with a well-developed herbaceous layer of smooth brome, mountain brome, fringed brome, Kentucky bluegrass and other Poa species along with some Idaho fescue. Shrubs other than common juniper included the two low-stature species, Sey's or prickly rose and creeping Oregon grape or low holly grape.

The first of these two slides presented another-- though somewhat "closer-to-it"--exterior view (following the two preceding "outside" views) while the second photograph showed the outer edge of this forest range as the photographer moved inside the forest. In the first photograph the taller trees were Douglas-fir except the tallest tree (center, behind Douglas-fir) was a ponderosa pine. There was a pole-sized ponderosa pine with a reddish-brown trunk between two Douglas-fir to right of big ponderosa pine and a ponderosa pine sapling in left-center foreground. Beside (immediately to right of) this ponderosa pine sapling there was quaking aspen. Somewhat larger than the ponderosa pine sapling in foreground were saplings of white fir. Grassy opening in right foreground of this first slide was dominated by smooth brome and Kentucky bluegrass, but immediately behind this native grasses dominated the herbaceous layer.

The second of these photographs featured a plant of common juniper (center foreground) and limber pine (three white-barked trunks in left foreground). Large seedlings to small saplings in foreground were of both Douglas-fir and white fir. All big trees (biggest trunk and one to its left-rear) were Douglas-fir. The well-developed and diverse herbaceous understorey was mostly smooth brome with some Kentucky bluegrass in foreground, but grasses behind that were less phenologically advanced included mountain brome, fringed brome, and Idaho fescue. Present throughout this herbaceous layer though at remarkably low density was cheatgrass.

North La Veta Pass (9400 feet), Costillo County, Colorado. Late June (estival aspect). FRES No. 21 (Ponderosa Pine Ecosystem). Mapped as Kuchler- 17 (Ponderosa Pine-Douglas-fir Forest). Some inconsistency here because Kuchler unit 17 is the only unit that that includes both ponderosa pine and Douglas-fir, but K-17 was listed under Ponderosa Pine Forest Ecosystem not Douglas-Fir forest Ecosystem. SAF 210 (Interior Douglas-Fir). White Fir-Douglas-Fir-Ponderosa Pine Series under Upper Montane Coniferous Forest (Dick-Peddie, 1993, ps. 62-64). Rocky Mountain Montane Conifer Forest 122.6, Douglas Fir-White Fir (Mixed Conifer Series) 122.61, Pseudotsuga menziesii-Mixed Conifer Association, 122.613 in Brown et al. 91993, p. 37). Southern Rocky Mountains- Sedimentary Subalpine Forest Ecoregion, 21b (Chapman et al., 2006).

165. Inside a Rocky Mountain mixed conifer forest- Interior of a upper montane mixed conifer forest range. Major tree species included Douglas-fir, ponderosa pine, white fir with common juniper, limber pine, and quaking aspen as associates. Shrubs were limited with Sey's or prickly rose and low Oregon grape being the two most abundant species in this forest. Deep inside this diverse forest major herbaceous species included mountain and fringed bromegrasses, Idaho fescue, and unidentifiable (at early phenological stages) Poa species. Smooth brome and Kentucky bluegrass were present but secondarily to the native perennial grasseses in the forest interior. Cheatgrass was present but not common.

There were a few forbs, but none were blooming at time of photographs. One of the most abundant forbs (if not the most abundant) was some Astragalus species whose compound leaves were conspicuous in both of these slides. .

Observation: the mixed conifer type of the Sierra Nevada Mountain has been widely recongized and commonly described as, for example, by the Society of American Foresters (Eyre, 1980). Surprisingly, the mixed conifer type of the Rocky Mountains has yet to receive the equivalent recognition even though it clearly exist as a distinctive forest cover type. For instance, this forest community is found throughout forested areas of New Mexico for which it was titled and described as mixed conifer forest by Dick-Peddie (1993, ps. 58, 62-64), New Mexico's reigning student of natural vegetation. Identification and description of forest cover types (Eyre, 1980) and rangeland cover types (Shiflet, 1993) seems as much as anything else in finding authors who will make the effort to write them up. This is probably even more the case for smaller or lesser known cover types.

North La Veta Pass (9400 feet), Costillo County, Colorado. Late June (estival aspect). FRES No. 21 (Ponderosa Pine Ecosystem). Mapped as Kuchler- 17 (Ponderosa Pine-Douglas-fir Forest). Some inconsistency here because Kuchler unit 17 is the only unit that that includes both ponderosa pine and Douglas-fir, but K-17 was listed under Ponderosa Pine Forest Ecosystem not Douglas-Fir forest Ecosystem. SAF 210 (Interior Douglas-Fir). White Fir-Douglas-Fir-Ponderosa Pine Series under Upper Montane Coniferous Forest (Dick-Peddie, 1993, ps. 62-64). Rocky Mountain Montane Conifer Forest 122.6, Douglas Fir-White Fir (Mixed Conifer Series) 122.61, Pseudotsuga menziesii-Mixed Conifer Association, 122.613 in Brown et al. 91993, p. 37). Southern Rocky Mountains- Sedimentary Subalpine Forest Ecoregion, 21b (Chapman et al., 2006).

166. Starting a new forest- A small clearcut of the white fir-Douglas-fir--ponderosa pne mixed conifer type on a ridgeline just behind the second-growth forest shown and described immediately above. The young conifers were pretty much an even mixture of the three climax dominants of Douglas-fir, ponderosa pine, and white fir. The most common and abundant major woody species was Gambel oak (Quercus gambelli). Gambel oak scrubland with associate species of common or alderleaf mountain mahogany (Cercorcarpus mopntanus) and New Mexico locusts (Robinia neomexicana) occurred contiguous with lower elevations of this same tract of forest.

North La Veta Pass (9400 feet), Costillo County, Colorado. Late June (estival aspect). FRES No. 21 (Ponderosa Pine Ecosystem). Mapped as Kuchler- 17 (Ponderosa Pine-Douglas-fir Forest). Some inconsistency here because Kuchler unit 17 is the only unit that that includes both ponderosa pine and Douglas-fir, but K-17 was listed under Ponderosa Pine Forest Ecosystem not Douglas-Fir forest Ecosystem. SAF 210 (Interior Douglas-Fir). White Fir-Douglas-Fir-Ponderosa Pine Series under Upper Montane Coniferous Forest (Dick-Peddie, 1993, ps. 62-64). Rocky Mountain Montane Conifer Forest 122.6, Douglas Fir-White Fir (Mixed Conifer Series) 122.61, Pseudotsuga menziesii-Mixed Conifer Association, 122.613 in Brown et al. 91993, p. 37). Southern Rocky Mountains- Sedimentary Subalpine Forest Ecoregion, 21b (Chapman et al., 2006).

167. Growing low- Holly grape, low holly grape, or low Oregon grape (Mahonia repens) on floor of a mixed montane forest. This interesting plant is a member of the Berberidaceae.

North La Veta Pass, Huerfanoco County Colorado. Late June, peak bloom stage of phenology.

168. A passing rose- Prickly rose (Rosa acicularis subsp sayi) growing in a small gap on on floor of a mixed montane forest. On this particular forest range there were just a few isolated plants of this species that were nowhere abundant enough to provide much by way of browse. Its presence did serve as an indicator of just how botanically rich this forest vegetation was.

North La Veta Pass, Huerfanoco County Colorado. Late June, peak bloom stage of phenology.

169. Rocky Mountain lodgepole pine forest- An exterior view of the Rocky Mountain form of the lodgepole pine range type showing details in structure of vegetation and plants (eg. retention of dead lower limbs on bole). This is the Rocky Mountain lodgepole pine (Pinus contorta var.latifolia). There was one prominent layer of woody plants (mostly shrubs; some, but rare, younger pines) and a less prominent layer of herbaceous species. The woody understorey was dominated by common juniper (Juniperus communis) with dwarf or broom huckleberry (Vaccinum scoparium) and myrtle huckleberry or blueberry (V. myrtillus subsp. oerophilum) as associate shrubs. Most common forbs were heart-shaped arnica (Arnica cordifolia) and one-sided wintergreen (Orthilia secunda). Grasses were absent, but there were a few "runt" individuals of Carex spp.

Apache National Forest, Grand County, Colorado. June. FRES No. 26 (Lodgepole Pine Forest Ecosystem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

170. Rocky Mountain lodgepole pine forest- Interior veiw of the Rocky Mountain form of the lodgepole pine range (forest cover) type. There was a prominent understorey consisting of a woody (mostly shrub) layer and, less prominently, an herbaceous layer. Dominant shrub was common juniper with dwarf or broom huckleberry and myrtle huckleberry of secondary importance. The herbaceous layer (or understorey if one cares to view layer and understorey as synonymous) was mostly forbs with heart-leaved arnica and one-sided wintergreen most common. For practical purposes there were no grasses and very few stunted Carex plants.

Apache National Forest, Grand County, Colorado. June. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

174. Lower elevational limit- Overall view of the lodgepole pine forest zone at its approximate lowest elevation (about 7500 to 8000 feet) in this area. This shot into the sun on a bright summer morning was a southeast slope. Elevational zones of ponderosa pine and Douglas-fir forest types were below.

Conspicuous shrubs were primarily antelope bitterbursh (Purshia tridentata) with some wax currant (Ribes cereum) and lesser cover of thimbleberry (Rubus parviflora). The dominant herbaceous species was timberline or glaucus bluegrass (Poa glauca subsp. rupicola= P. rupicola).

Details of the interior of the lodgepole pine forest range type were shown below.

Roosevelt National Forest, Larimer County, Colorado. Early July. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). There should be-- but was not--a Pinus contorta Association and/or Series within (under) the Rocky Mountain Montane Conifer Forest 122.6 in Brown et al. (1998, p. 37). How did they miss this one? Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

175. Exterior of a lodgepole pine forest range- Physiogonomy and structure of the Rocky Mountain lodgepole pine range type. An uneven-aged population of lodgepole pine that was at the state of old-growth (though most likely not a virgin forest by any means). This was clearly climax vegetation from standards of species composition and, except for more old and senescing pines, structure. Pine reproduction was limited, but adequate for maintenance of the population. The understorey consisted of at least one shrub layer, two herbaceous layers, and one thallophyte layer (on granite rocks). These layers were shown and described immediately below.

This lodgepole pine forest range was on a general rock outcrop-granitic soil forest site (south to southeast slope) just a few hundred feet above the lower elevational limit of this forest cover type in the Southern Rocky Mountains.

Roosevelt National Forest, Larimer County, Colorado. Early July. Elevation was approximately 8000 feet. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). There should be-- but was not--a Pinus contorta Association and/or Series within (under) the Rocky Mountain Montane Conifer Forest 122.6 in Brown et al. (1998, p. 37). How did they miss this one? Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

176. Now for the interior- Structure and species composition of a Rocky Mountain lodgepole range on a granite rock outcrop in the Southern Rockies. This was an uneven-aged (seedling to senescing adults) population of lodgepole pine with an understorey of at least one shrub layer, two herbaceous layers, and one thallophyte layer (on granite rocks). The major shrub species were antelope bitterbrush, common juniper, bearberry manzanita or kinnikinnick (Arctostaphylos uva-ursi) with dominance and associate rank varying at local scale. There was also cover of wax currant, but this was much less than that of the preceding three species. The dominant herbaceous species was glaucus bluegrass with lanceleaf stonecrop (Sedum lanceolatum), sulphur-flowered wild-buckwheat (Erigonum umbellatum), and sagewort wormwood or field sagewort (Artemisia campestris subsp. caudata) the main forb species.

The darker green mat in center foreground was bearberry manzanita or kinnikinnick. This member of the Ericaceae was also shown in the next two slides which indicated just how abundant this member of the heath family was in the lodgepole forest cover type in this general area.

Roosevelt National Forest, Larimer County, Colorado. Early July. Elevation was approximately 8000 feet. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). There should be-- but was not--a Pinus contorta Association and/or Series within (under) the Rocky Mountain Montane Conifer Forest 122.6 in Brown et al. (1998, p. 37). How did they miss this one? Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

177. More views on the inside- Two sequential (moving closer-in) shots of the interior of a lodgepole pine forest range on a typical granitic soil-rock outcrop site (south to southeast slope) in the Southern Rocky Mountains.

Most of the darker (or birghter)- green "plant carpet" was bearberry manzanita or kinnikinnick. A large colony of this mat-forming ericaceous shrub as was also shown in the preceding slide.Numerous plants and substantial cover of antelope bitterbrush and common juniper as were cespitose individuals of glaucus bluegrass were also visible in these two photographs. Seedlings, saplings, and small poles of lodgepole pine were distinctly visible in both photographs.

On this forest site of relatively low production potential, lodgepole pine had developed into an uneven-aged forest with comparatively low tree density (low relative density; a relatively sparse cover of lodgepole pine). Reproduction and establishment of interior lodgepole pine was discussed below in the caption beneath two small (young) yet cone-bearing saplings.

Roosevelt National Forest, Larimer County, Colorado. Early July. Elevation was approximately 8000 feet. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). There should be-- but was not--a Pinus contorta Association and/or Series within (under) the Rocky Mountain Montane Conifer Forest 122.6 in Brown et al. (1998, p. 37). How did they miss this one? Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

178. Plenty of rock- Outcrops of granite adorned the floor of a lodgepole pine forest range in the Southern Rocky Mountains. This was an uneven-aged population of lodgepole pine at or approaching old-growth status with limited (though apparently adequate) pine reproduction. On the shallow soil of this south to southeast slope the adult pines seen here were nearing (or at) their maximum size. Some of the pines in this forest were at the senescing stage and seedlings and saplings were clearly visible.

The dominant (most abundant) forb in this view was lanceleaf stonecrop with glaucus bluegrass, sulphur-flowered wild-buckwheat, and sagewort wormwood or field sagewort also important. Various lichens had colonized the outcropping granite. This thalophytic component was shown in the next set of slides.

Roosevelt National Forest, Larimer County, Colorado. Early July. Elevation was approximately 8000 feet. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). There should be-- but was not--a Pinus contorta Association and/or Series within (under) the Rocky Mountain Montane Conifer Forest 122.6 in Brown et al. (1998, p. 37). How did they miss this one? Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

180. Slightly different site, but same story- Lodgepole pine forest range on a predominately south slope in the Southern Rocky Mountains. This montane forest vegetation was roughly 500 feet higher in elevation than that featured immediately above. This soil profile was also noticeably deeper, but there was still more than enough granite rock. Like the preceding example, this was most likely climax--certainly, subclimax--vegetation. All of the larger pines were young adults. Though these lodgepole pine were approaching the size of those in the preceding example of this forest cover type the example forest featured here was on a different forest site that had greater potential for tree growth (even though degree of land slope was drastically greater in this example). Thus final adult tree size would be considerably larger on this potentially more productive site (see next slide).

In contrast to the uneven-aged population of lodgeplole pine in the example of this range type presented immediately, the population of this example was even-aged lodgepole pine.Correspondingly, tree density and canopy cover was considerably greater in this lodgepole pine forest. Density and size of individual plants of glaucus bluegrass was also greater, but relative cover of antelope bitterbrush and, especially, of most forb species was much less. There were some scattered plants of wax current (Ribes cereum). (These vegetational features were more obvious in the next two slides.)

On this forest range, forb species were more limited than on the shallower and rockier soil of the preceding example. The dominant forb on this edaphically more favorable site was field sagewort or sagewort wormwood (Artemisia campestris subsp. caudata), but the overall dominant herbaceous species was glaucus bluegrass (and "hale and hearty" specimens at that). Glaucus bluegrass had to share understorey dominance with antelope bitterbrush. This was a sho' 'nough productive forest range with both abundant forage (herbage) and browse crops. Prairie or fringed sagewort (Artemisia frigida) joined its fellow genus member, sagewort wormwood or field sagewort, noted above in the different form of lodgepole pine range vegetation treated here.

Roosevelt National Forest, Larimer County, Colorado. Early July. Elevation was approximately 8500 feet. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). There should be-- but was not--a Pinus contorta Association and/or Series within (under) the Rocky Mountain Montane Conifer Forest 122.6 in Brown et al. (1998, p. 37). How did they miss this one? Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

181. More tree cover, more pine duff, and more bluegrass- An even-aged lodgepole pine forest range with relative high tree density and crown cover in the Southern Rocky Mountains. On more open portions of this forest (including both small parks and local stands of wider tree spacings) the understorey included both a shrub and an herbaceous layer. This was shown in the immediately preceding two photographs. Where tree density and canopy cover were greater (as shown in these two slides) the understorey was mostly limited to a sporadic (interrupted) herbaceous layer comprised almost exclusively of glaucus bluegrass as shown in these two photographs. This was also in contrast to the multilayered understorey on the lodgepole pine forest range detailed in the previous example of lodgepole pine range on a forest site of lower production potential.

Attention was also drawn to the conspicuously greater cover of pine straw (the ground cover layer composed of shed pine needles) on this forest of even-aged trees of higher denser and canopy cover. What, if any, impact greater cover of pine needles had was not known.

Larimer County, Colorado. Early July. Elevation was approximately 8500 feet. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). There should be-- but was not--a Pinus contorta Association and/or Series within (under) the Rocky Mountain Montane Conifer Forest 122.6 in Brown et al. (1998, p. 37). How did they miss this one? Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

182. The very old and large, the very young and small, and those in between- An uneven-aged stand of lodgepole pine on an east slope in the Southern Rocky Mountains. Two old-growth specimens with two seedlings of two sizes at their base and saplings to small poles behind pretty much covered the range in age/size classes. Only tiny new seedlings were missing.

"Lodgepole pine is very intolerant of shade and competition from other plant species" (Burns and Honkala, 1990) with seedlings and other smaller trees typically not thriving under shade. Viewers can see that there was mostly "open sky" in the scene shown here so that younger trees were not subjected to much shade regardless of their age/size classes. In more open forest, especially with widely scttered adult trees, younger lodgepole pines can establish and mature so that uneven-aged lodgepole pine forests sometimes develop. Such was the situation shown here and in the example before the last one above.

The vegetation seen here was a different local forest (and perhaps different forest site) from the two immediately preceding examples. All of three forest locations were within a half-mile radius (or less) of each other. Thie forest shown here was private property in contrast to the public land of a national forest in preceding examples. The present example of lodgepole pine forest had been stocked heavily with horses several years earlier. There was almost no understorey in this lodgepole pine stand. Herbaceous plants were limited to almost exclusively forbs, mostly to white or pale locoweed (Oxytropis sericea) with a respectable showing by the native crucifer, hoary false alyssum (Barteroa incana). The photographer could not help wondering if any horses had been locoed on this wooded range. Maybe that was why it was destocked at time of photograph.

Bark color, patterns, etc. of the two old-growth pines were in stark contrast to each other with the farther tree having smooth bark and the nearer one having furrowed bark. There is considerable variation in bark features among lodgepole pines. Bark color ranges from reddish orange as in these two trees to gray. Bark on most adult lodgepole pine remains relatively smooth, but on some boles the bark is furrowed (as in the foremost trunk here) much like that of ponderosa pine (a source of confusion where the two species occur together). The nearer of these two pines bore the battle scar from a run-in with lightning, an all-too-common atmospheric phenomenon in the Rocky Mountains.Lightning-ignited fires have been a natural part of range ecosystems since there was vegetation to serve as fuel.

Larimer County, Colorado. Early July. Elevation was approximately 8500 feet. FRES No. 26 (Lodgepole Pine Forest Ecosytem). K-8 (Lodgepole Pine-Subalpine Forest). SAF 218 (Lodgepole Pine). There should be-- but was not--a Pinus contorta Association and/or Series within (under) the Rocky Mountain Montane Conifer Forest 122.6 in Brown et al. (1998, p. 37). How did they miss this one? Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

183. Early producers- Lodgepole pine is one of the earliest sexually maturing Pinus species in western North America. This precocious species can produce viable seed at ages as young as 5 to 10 years (Harlow et al.,1979, p. 111; Burns and Honkala, 1990), literally baby trees having babies. These two pines were slightly less than five feet tall and both bore cones of ripening seed. Undoubtedly, early seed production is evolutionary adaptation to fire-prone environments (recall the lightning-struck adult lodgepole pine from the preceding photograph). Cones of Pinus contorta var. latifolia are serotinous meaning that cones are retained on trees without opening for more than one year after reaching maturity (Helms, 1998). The serotinous cones of inland or interior lodgepole pine usually open only when there is adequate heat accompanied with a dry atmosphere, conditions most commonly present only with fire or extremely hot weather.

Interior lodgepole pine forests comprise are one of the most pronounced of fire types (cover types adapted to fire and that burn frequently or readily). This commonly results in establishment of dense poplulations of pines of the same age (cohort). These "doghair stands" are often stocked with small, growth-stunted trees (ie. overstocked) which are more prone to crown fires. Natural establishment of even-aged, overstocked lodgepole pine forests is more apt to take place on forest sites that are more favorable for tree establishment rather than on marginal sites on which there can be forests having uneven-aged populations with lower tree density (recall an example of this that was presented above).

The two young cone-bearing lodgepole pines in this photograph were growing on the edge of the even-aged forest shown in the example (two two-slide/caption sets) before the preceding slide.

Roosevelt National Forest, Larimer County, Colorado. Early July.

184. Ripe and breaking off- Terminal portion of a leader of lodgepole pine with mature female (cones). The cones of Rocky Mountain lodgepole pine (Pinus contorta var. latifolia) most commonly break off above their bases so that the bottom part of cones (ie. some of the basal scales) remain attached to the cone-bearing branches. One such instance of such was presented in this photograph.

Lodgepole pine typically has two needles per fascicle. Most needles in this area were about three inches long, at the upper limit of leaf length for this species

Larimer County, Colorado. Early July.

185. Separate-sex cones- Male or pollen cones, microsporangiate strobili, (on two branch terminuses at two left) and female or seed cones, megasporangiate strobili (on branch terminus at right) of Rocky Mountain lodgepole pine.

The details of strobili in the gymnosperms was detailed above when discussing Douglas-fir.

Larimer County, Colorado. Early July.

186. Big girls and little girls- Megasporangiate strobili (female or seed-baearing cones) of Rocky Mountain lodgepole pine on a single leader. The first photograph presented mature cones at left or lower on the branch and immature (pollen-receptive) cones at right or at terminus of branch. The second slide was a view of the terminal portion of this branch so as to feature the immature megasporangiate strobili. This second photograph also showed the the two-needle fascicles of this species.

Larimer County, Colorado. Early July.

187. Now the boy's turn- Male or pollen cones (microsporangiate strobili) of Rocky Mountain lodgepole pine. Also fairly decent shot of needles of this species. This was an early morning ( shortly after sunup) shot after a nice shower the night before.

Larimer County, Colorado. Early July.

188. Grass neighbor- Glaucous or timberline bluegrass (Poa glauca subsp. rupicola= P. rupicola) was the dominant herbaceous species in the understorey of lodgepole pine forest of twn forest sites in the Southern Rocky Mountains. The photographs in this and the next two sets of slides were of glaucous bluegrass growing on ta south slope of relatively deep, granite-derived soil

The first of these two slides showed a larger (presumedly older) plant surrounded by several smaller (younger) plants of glaucous bluegrass along with some small plants of field sagewort (Artemisia campestris subsp. caudata). The second slide presented several still-yet larger plants of glaucous bluegrass with a plant of wax current (Ribes cereum) in the left background. All adult shoots of glaucous bluegrass were bearing panicles.

Roosevelt National Forest, Larimer County, Colorado. Early July.

189. Examples of timberline or glaucous bluegrass- Two large plants of glaucous (timberline) bluegrass at peak bloom served as nice examples of this species which was the dominant herbceous species in understorey of lodgepole pine forest range in the Southern Rocky Mountains. Most panicles of these plants were at immediate post-anthesis stage. Both also had a few sexual shoots from the previous year that remained intact except for shed spikelets. There were a few shoots of field sagewort in the first slide.

Timberline bluegrass is strictly cespitose (ie. tillers only; no stolons or rhizomes).

Roosevelt National Forest, Larimer County, Colorado. Early July.

190. Infloresceances of glaucous or timberline bluegrass- Several examples of panicles on glaucous bluegrass growing on a lodgepole pine forest range. The panicles of this species are relative large and generally described as narrow and somewhat tight or compressed.

Spikelets of this species usually have a conspicuous purple coloration when young, but they take on a pale amber color as they near maturity. Spikelets in three photographs were in milk grain stage and had amber rather than the distinctive purplish color.

Interesting (and practical) point: the Poa species are a "taxonomic challenge" even to the experienced agrostologist. Many bluegrass species closely resemble each other differing only in details some of which are minute or, even trivial. This situation certainly exist for P. glauca and related species. Barkworth et al. (2007, ps. 576-579) described close resemblance of P. glauca, especially P. glauca subsp. rupicola, with several other Poa species especially P. interior and P. nemoralis. According to these workers confusion among Poa taxa is more apt to occur among herbaria specimens with positive identifiction frequently depending on details of features like pubesence, veins, keels, and calluses of lemma and/or palea. P. glauca is extremely variable being "highly polyploid and presumed to be highly apomictic" (Brkworth et al., 2007, p. 577). As if that was not enough, P. glauca hybridizes with other Poa species including the widespread and versatile P. secunda.

Roosevelt National Forest, Larimer County, Colorado. Early July; milk stage of caryopses.

191. Small and showy- Lanceleaf or spearleaf stonecrop (Sedum lanceolatum) in understorey of a Rocky Mountain lodgepole forest range in the Southern Rocky Mountains on a granite rock outcrop forest site. This species was the most abundant forb in the herbaceous layer(s) of this range.

The Crassulacean Acid Metabolism (CAM) photosynthesis pathway was discovered in the Crassulaceae family of which Sedum is a major genus. The habitat of lodgepole forests of this form is a classic CAM-requiring environment: dry soils, short-growing season, frequently limited light, high elevation, severe competition. Light use efficiency is a necessity. Ergo, some of the most successful understorey plants in high elevtion, semiarid, lodgepole pine forest range are those species which evolved traits that use light more effectively and efficiently. Darwin's "survival of the fittest" where survival translates to successful reproduction (survival of the species not necessarily of an individual plant).

Anyway lanceleaf stonecrop is a common and very showy species even if it is a small plant.

Roosevelt National Forest, Larimer County, Colorado. Early July.

192. A memory refresher- Another specimen of common juniper (Juniperus communis). An example of this range conifer was presented earlier, but the one shown in this photograph was growing on the forest range described more recently (the granite rock outcrop forest site) and accompanied by sagewort wormwood or field sagewort (Artemisia campestris subsp. caudata).

Roosevelt National Forest, Larimer County, Colorado. Early July.

193. Colony of fireweed or willowherb (Epilobium augustifolium= Chamaenerion augustifolium)- This member of the evening primrose family (Onagraceae) got it's more common common name from it's adaptation to fire, especially intense forest fires. Fireweed is a pioneer species-- usually the most prominent pioneer-- following fires. This perennial species is found in various regions of North America, but is most widely distributed in the general Rocky Mountain and Pacific Slope Regions.

Fireweed is especially valuable in watershed stability and protection following fires that denude steeply sloping land. It has been regarded as being of fair to good forage (when immature) for small ruminants. The showy, bright-pink inflorescences and graceful leaves impart aesthetic values to this forest and range forb as well.

Fireweed was one of the 200 range plant species included on the Master List for the International Range Plant Identification Contest sponsored by the Society for Range Management. The various editions of North American Range Plants (Stubbendieck et al., 1981, 1982, 1992, etc.) provided complete, yet concise, descriptions of E. augustifolium. Readers were referred to those excellent works. This species was also included in the Range Plant Handbook (Forest Service, 1941, p. W50) and Notes on Western Range Forbs (Hermann, 1966, ps. 185-186).

This robust clump was photographed in the understory of a mature lodgepole pine forest (elevation about 6000 feet) in Yellowstone National Park (and even before the 1988 holocaust). While fireweed is a pioneer and rapidly invades burnt-over ground it still persist into the climax forest as shown in this slide. July.

195. Death cometh on wings- Mixed conifer (lodgepole pine, ponderosa pine, and Douglas-fir, were major conifers; there were a few isolated Engelmann spruce) montane forest in Never Summer Mountains. The forest community shown here was on the Continental Divide or Great Divide at an elevation of about 10,640 feet on a predominately south and west aspect. At this elevation forests of Engelmann spruce and subalpine fir are the typical natural plant community, but at this particular location, and on predominately south and southwestrn slopes, the subalpine Picea-Abies forest cover type developed at higher elevations resulting in a mixed conifer transition forest (= a forest ecotone) between subalpine and montane zone coniferous forests.

The most striking phenomenon (and obvious lesson) from this pair of photographs was death of almost all pines (and the few Engelmann spruce) by feeding of a beetle known variously as mountain pine beetle, pinebark beetles, Black Hills beetle, or Rocky Mountain pine beetle, (Dendroctonus ponderosae). This is a weevil native to North America and its role as a natural herbivore and limiting factor in Pinus populations has been remarked for years, as for instance by Shelford (1963, p. 179). Remaining live conifers in these two photographs were Douglas-fir. There was an almost complete kill of the two pine species and Engelmann spruce. The latter were preyed on by the spruce beetle (Dendroctonus rufipennis). (More of outbreak of the spruce beetle below.)

Herbaceous species were very sparse in the current forest with grasses (mostly Poa and Bromus species) and a few sedges (Carex spp.) making up an interrupted herbaceous layer. These gramnoids could be expected to be released by loss of the former dense canopy of tree crowns.

The most noticable and, more importantly, both the ecological and economic aspects of mountain pine beetle herbivory is the killing of trees in patches--often by the hundreds of acres--rather than as isolated, individual trees. Tree death generally is due to mass colonization and feeding of many weevil larvae inside of their egg and, later, feeding galleries. The female beetle lays groups of eggs inside a gallery she made. Upon hatching the larvae feed on wood in larval galleries mined at right angles to their mother's egg gallery. This mining-feeding activity extends into the phloem tissue and destroys the cambium (at least cambial function) ultimately causing tree death. Trees respond to beetle activity by producing resins to block the galleries. Some species like D. ponderosae are also vectors for the bluestain fungus (Grosmannia clavigera). This saprophytic fungus functions together with mining/feeding action of mountain pine beetle to eventually kill the tree. This ecological relationship of prey tree and insect/pathogen is complex and incompletely understood even after much research (Graham, 1967, Paine et al., 1997). The life cycle of D. ponderosae is one year (one generation annually) in most of the Rocky Mountain System.

Interested readers can find much useful information about the various pine beetle species from a number of sources ranging from standard texts in Forest Entomology, including the standard by Graham and Knight (1965), to reputable, on-line Agricultural Extension publications, including some from Colorado State University and University of Idaho, and materials put out by the US Forest Service. Good sources for the ecological relationship between beetles and fungus include Mitton and Sturgeon (1982) and Schowalter and Filip (1993).

Aggregation pheromones released by adult pinebark beetles attract more adults and soon a tree is under "full attack". Trees are more apt to be attacked under certain conditions including stress due to drought or crowding which is much more likely to occur in "doghair" stands of pine (which, coincidentially, could have been alleviated by natural thinning under low-intensity natural fire or proper prescribed burning.

The central interest from the perspective of the forest plant community and, at larger scale, of forest range ecosystem and landscape is the roles or impacts--both short-term and long-term--of large, severe outbreaks (epidemics) of mountain pine beetle. This native forest insect has influenced forest ecosystems for millenia and in that regard amounts to natural disturbance the same as drought, flood, fire, temperture extremes, avalanches, wind and ice storms, as well as herbivory by other native animals. If conifers like lodgepole and ponderosa pine, Engelmann spruce, and subalpine fir could not co-exist with native insects and pathogen like mountain pine beetle and blue stain fungus these tree species (and their pests) would have been extinct ages ago.

There are almost countless scientific publications devoted to bark beetles and their influence on forests. A readily read synopsis of lodgepole pine and moutain pine beetle included that of Kaufmann et al (2008) which also included an anlysis of the interaction of fire with beetle/fungus-induced mortality. Studies in peer-reviewed periodicals included impact of mountin pine beetle on lodgepole forest succession (Amman, 1976 and regulation of pirmary productivity (Romme et al., 1986).

Nevr Summer Mountains (10,640 feet elevation), Rocky Mountain National Park, Grand County, Colorado. Mid-June (late vernal aspect). FRES No. 21 (Ponderosa Pine Ecosystem). Mapped as Kuchler- 17 (Ponderosa Pine-Douglas-fir Forest). SAF 237 (Interior Ponderosa Pine); no SRM for this pine type per se, but it would be the Southern Rocky Mountain equivalent of SRM 109 or 109 and 110 combination (Ponderosa Pine-Shrubland-Grassland). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

196. Yikes, we're being invaded!- A seral stand of young quaking aspen (Populus tremuloides) being invaded by Engelmann spruce and subalpine fir. This drama of Frederic Clements "dynamic vegetation" was on the boundary between upper montane and subalpine zones. Sucessional replacement of the "quakies" by spruce and spruce is the fate of seral aspen communities at elevations of subalpine forests Beidleman et al., 2000, ps. 14-15).

This process was probably accelerated by extremely heavy browsing by Rocky Mountain elk (Cervus canadensis subsp. nelsoni) which was obviously stressing some of the aspen shoots as was apparent in the second of these two photographs (this second slide was a closer-in version of foreground in the first slide).

Kashian et al. (2007) evaluated dynamics of quaking aspen stands in northcentral Colorado and reported that dominance by various conifers was increasing on over one-third of aspen stands studied. About 15 percent of quaking aspen stands in this area were declining with decline most severe under heavy browsing by elk and/or fire exclusion. Kaye et al. (2005) analyzed impacts of elk browsing and natural succession (invasion) of native conifer species on quaking aspen in Rocky Mountain National Park and adjacent areas. Findings of Kaye et al. (2005) indicated that elk browsing and conifer invasion decreased aspen regeneration (establishment), but did not affect growth rate of established trees (clonal shoots). From studies like these it would be concluded that management to increase quaking aspen abundance should be focused on reproduction (recruitment) of new aspen shoots or, probably less commonly, trees from seed. Otherwise, in absence of natural or prescribed fire and/or with continued heavy browse use by elk, coniferous species like Engelmann spruce and subalpine will continue to replace quaking aspen, at least on those forest sites for which conifers are the potential natural vegetation in absence of fire, outbreaks of bark beetle (Dendroctonus spp.), avlanches, etc.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect).FRES No.19 (Aspen-Birch Ecosystem). SRM 411 (Aspen Woodland). SAF 217 (Aspen). Populus tremuloides subclimax Association in Douglas Fir-White Fir (Mixed Conifer) Forest Series of Brown et al.(1998, p. 37). Succession is taking this to: FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Enlgemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

197. Transition zone forest between Douglas fir on north slope and subalpine fir (locally dominant) and Engelmann spruce-subalpine forest cover types.

Rocky Mountain National Park, Larimer County, Colorado. August. FRES No. 20 (Douglas-fir Ecosystem) and FRES No. 23 (Fir-Spruce Ecosystem) with respective Kuchler units K-11 (Douglas-fir Forest) and K- 19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir) and SAF 210 (Interior Douglas-fir) transition. Ecotone between Englemann Spruce-Alpine Fir Series and Douglas Fir-White Fir (Mixed Conifer) Series of Brown et al. (1998). Southern Rockies-Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

198. Landscape-scale view of Englemann spruce-subalpine fir forest- Appearance of this forest range cover type in early summer at the northern end of the Front Range of the Southern Rocky Mountains. This landscape photograph included the Subalpine and the Alpine life zone as generally used today. In the original life zone scheme (of seven zones) devised by C. Hart Merriam there was no Subalpine. Instead what is commonly called Subalpine in current usage was part or all of Merriam's Canadian and Hudsonian Zones. The Alpine, defined as being above timberline, is the same in both Merriam's and contemporary schemes.

There have also been differences in the use of Southern Rocky Mountains and Central or Middle Rocky Mountains. Fenneman (1931, ps. 92-182 passim) delineated the Southern Rocky Mountain physiographic province as extending through the Laramie Mountains of Wyoming while the Middle Rocky Mountain province included the Yellowstone Plateau, Bighorn Mountains, Tetons, Wasatch Mountains, and Unita Mountains, among others. Shelford (1963, ps. 160-181) divided the Central Rocky Mountains and the Southern Rocky Mountains differently from this. He included the Colorado Rockys as being in the Central division along with the Unita and Wasatch Ranges (Shelford, 1963, p. 160).

Either way this slide showed appearance of the Englemann spruce-subalpine forest cover type growing on the generic subalpine mountain zone inside the Front Range.

Larimer County, Colorado. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998).Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

199. Englemann spruce-subalpine fir forest- An exterior view of a stand of old-growth Englemann spruce- subalpine fir showing the physiogonomy and structure of this climax forest type. The Society of Amreican Foresters (Wenger, 1984, p. 3) ranked Englemann spruce as Tolerant and subalpine fir as Very Tolerant. The ecological veracity of these ratings was proven in this photograph. Both of these climax dominants were "reproducing in their own shade". The three distinctively green trees of shorter stature in the foreground were young Englemann spruce. Most of the shortest, smallest conifers with the distinctive bluish-green needles were Colorado blue spruce (Picea pungens), a species with tolerance rated as Intermediate. These blue spruce along with some subalpine fir were growing in openings recently created by fallen subalpine fir. Similar colored conifers in the midground between the foreground young trees and the old-growth conifers of the background were subalpine fir of somewhat older age and larger size.

The sparse understorey comprised primarily of Vaccinium species and was not shown in these exterior "shots" of this forest cover type.

Apache National Forest, Grand County, Colorado. June. FRES No.23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

200. Exterior view of a climax (and old-growth) Englemann spruce and subalpine fir forest in the Front Range of the Southern Rocky Mountains- Composite "shot" of a magnificant Rocky Mountain subalpine forest with a "picture-perfect" display of the species composition. The immense conifer on the far right margin was a subalpine fir as was the younger tree with the conspicuous grayish bark immediately to it's left and the smaller one to it's left. Note that the Very Tolerant subalpine fir were growing under shade of the Englemann spruce (the tall trees in the center and right-center background with the orangish- or reddish-brown trunks). Eyre (1980, p. 86) observed that while these two conifers were generally codominants there were areas and forest sites on which "subalpine fir is the major climax species" while Englemann spruce"is a persistent long-lived seral species". That was at least the partial apparent pattern shown here. Note also crown shape and dead limbs of the Englemann spruce patriarchs. Younger conifers in the left foreground were Englemann spruce indicating again successful regeneration of this Tolerant species. Regeneration of these two species beneath what were obviously old-growth individuals attested to the climax nature of this vegetation.

Understorey composition was primarily Vaccinium and Ribes species, some of which were shown below in this series of slides. Individual plants of Poa and Carex species were present in trace amounts.

Apache National Forest, Grand County, Colorado. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

201. "Closer in" view of the right side of the preceding photograph- Regeneration of both Englemann spruce (larger of the saplings in right-of-center foreground and in left foreground) and subalpine fir (smallest foreground sapling to immediate left of the "big momma" parent subalpine fir.

Apache National Forest, Grand County, Colorado. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

202. Colorado blue spruce in a sunlite opening in Englemann spruce-subalpine fir forest- This blue spruce was growing in an opening formed by a fallen subalpine fir in the forest in the three preceding photographs. Blue spruce is rated as Intermediate in tolerance. This species could regenerate in this climax forest only in microhabitats having more light than in the typical understorey environment of this old-growth forest.

Apache National Forest, Grand County, Colorado. June.

207. Whitestem, smooth or wine gooseberry (Ribes inerme)- Various Ribes species are found thoroughout the forests and adjoining general habitats of North America. Ribes is particularily well represented among the western coniferous forests. Dayton (1931, p. 40) reported that the widely distributed R. inerme displayed here was one of the most common, best known of the western gooseberries. Gooseberry and current are common names for the same or two closely related genera. Some authorities distinguished those plants of common name, gooseberry, as Grossularia species whereas those designated as currents were presented as Ribes species. Some authors have relied on as many as four genera for these plants. Currently, most authors lump all into Ribes. The common distinction between goosebery and current is that the latter bear few or no spines or prickles on their stems. More consistently, currents have jointed pedicels such that their fruit breaks off cleanly from this "stem" whereas the fruit of gooseberries remains attached to this pedicel thus necessitating "snipping" prior to making jams or pies (to which avid gooseberry pickers like your author can redily attest).

The nearly countless publications devoted to native plants (ranging from florae or manuals to field guides) have usually given good descriptions of these interesting shrubs. For their relevance and application to Forestry and Range Management the following classic treatments were recommended: Dayton (1931, p. 40-43) and Forest Service (1940, p. B130-B 133).

208. Flowering leader of whitestem, wine, or smooth gooseberry- This specimen grew adjacent to an Englemann spruce near Larimer Pass. The fruit of Ribes is a berry. It is often a valuable feed source for wildlife species including birds and rodents. Browse from Ribes species is of fair feed value for native ruminants.

An unfortunate turn of events combined with biological relationships led to reduction of Ribes species in regions renowned for the highly valued white pine species (Haploxylon subgenus of Pinus). Ribes serves as intermediate host for the accidentlly introduced white pine blister rust (Cronartium ribicola). In efforts to save the white or soft pines, especially western white pine (Pinus monticola), widespread Ribes eradication programs were conducted in efforts to control or, hopefully, eradicate the white pine blister rust. The effort was unsuccessful and preservation of white pines depends on breeding for rust-resistant genotypes. Fortunately Ribes species were not exterminated in an understandable but misdirected management practice. Ribes species suffer only minor damage when infected by C. rubicola.

Near Larimer Passs, Jackson County, Colorado. June.

209. Interior of a stand of old-growth Englemann spruce- The understorey of a virgin Englemann spruce forest with an exclusive ground layer of red-fruit gooseberry, false gooseberry, or cloudcap current (Ribes montigenum). Several of the Ribes species have "gobs" of common names. This species has proven to be somewhat confusing in that that is regarded as being in the Ribes group known as currents and, despite presence of prickles or spines, not in the gooseberry group that at times has been designated as the separate genus of Gossularia.

Readers should note regeneration of Englemann spruce, a species rated as Tolerant (Wenger, 1984, p. 3). This is an example of a "pure stand" of Englemann spruce. Eyre (1980, p. 86), citing other workers, specified that in the Englemann spruce-subalpine fir forest "pure stands of either species can be found". Such a stand was shown here.

Rocky Mountain National Park, Larimer County, Colorado. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Enlgemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

210. Englemann spruce-subalpine fir forest on shallow southeast slope- A marginal site for the Englemann spruce-subalpine fir cover type that is in a transition to a limber pine (Pinus flexilis) forest. This shallow site was an example of one kind of a subalpine forest in the Front Range of the Rocky Mountains just below timberline. Fir and spruce trees lacked the massive size of those in slides shown above on sites that were more favorable for development of the fir-spruce subalpine forest. Individuals of limber pine were abundant in this vegetation that was adjacent to a limber pine forest that was presented below. This marginal, transitional forest (ie. combination forest primarily of the Englemann spruce-subalpine fir cover type with components of the limber pine type) had a better developed and more species-rich understorey than many of the subalpine forests in the Southern Rocky Mountains. It was an example of of a Rocky Mountain subalpine forest range of the sort valuable for both livestock and wildlife, especially big game like elk and mule deer.

Rocky Mountain National Park, Larimer County, Colorado. Elevation roughly 10,000 feet. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir), but with scattered limber pines and conterminous with an old-growth stand of limber pine (SAF 219). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

211. Understorey of Englemann spruce-subalpine fir on a shallow, rocky, southeast slope- Detail of the understorey of the subalpine transition forest shown immediately above. The understorey dominant was grouseberry or broom huckleberry (Vaccinium scoparium). Some species of bluegrass was (or where) present, but none had bloomed or set seed so the author could not identify it (them). One of the more common bluegrasses on such habitat is inland bluegrass (Poa interior). Muttongrass (P. fendleriana) is often locally dominant. The two large clumps of grass were the alien but naturalized orchardgrass (Dactylis glomerata) that served as yet another illustration as how pervasive many of the introduced or agronomic species have become through naturalization. The orchardgrass plant on the left was mostly hidden behind a clump of some species of Carex. Beidleman et al. (2000) described over 60 Carex species growing in Rocky Mountain National Park! Cones of both limber pine and Englemann spruce were visible in the photograph.

Rocky Mountain National Park, Larimer County, Colorado. Elevation about 10,000 feet. June. FRES No. 23 (Fir-Spruce Forest Ecosystem). K-14 (Western Spruce-Fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir).

212. Population of grouseberry, grouse whortleberry, small-leaved huckleberry, dwarf huckleberry, or broom huckleberry (Vaccinium scoparium)- Members of the heath family are some of the most common understorey plants in mountain forests of western North America. Several Vaccinium species are understorey dominants or associates of the Englemann spruce-subalpine fir forest range cover type. V. scoparium is one of these that is valuable as browse for big game animals, sheep, and cattle as well as for it's fruit which is used by small animals like rodents and birds, uncluding upland game birds. Dayton (1931, p. 135) described this species as "... perhaps the most abundant and widespread as well as the smallest and smallest leaved of the western (so-called) huckleberry species".

This sample was in the understorey of the subalpine transition Englemann spruce-subalpine fir forest shown in the preceding two slides (a "sub-sample" of the understorey displayed in the preceding slide).

Rocky Mountain National Park, Larimer County, Colorado. Elevation about 10,000 feet. June.

215. Something to grouse about- The ground or lowest layer (also the third woody layer) of an Engelmann spruce-lodgepole pine-quaking aspen-sublpine forest. This ground or lower shrub layer consisted almost exclusively of Vaccinium scoparium which is known variously as grouseberry or grouse whortleberry or, perhaps as more commonly known among foresters and rangemen, broom, dwarf, or littleleaf huckleberry.

Second photograph showed dung of elk which were heavily browsing the dwarf huckleberry. Dunging is one aspect or category of herbivory. Also presented in this second slide were cones of Engelmann spruce and lodgepole pine.

Rocky Mountain National Park, Grand County, Colorado. Mid-June (late vernal aspect). FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

216. Engelmann spruce- subalpine fir forest range— Rocky Mountain subalpine forest; petran subalpine forest association of Clements. Engelmann spruce (red trunks, foreground); subalpine fir (white bark, background); conspicuous understory forb iswooly ragwort (Senecio atratus).Elevation about 11,000 foot level. Hudsonian life zone of C. Hart Merriam.

Rocky Mountain National Park, Larimer County, Colorado. August, estival aspect. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

217. Engelmann spruce-subalpine fir forest range type— At an elevation of approximately 11,000 this is less than 1000 feet from timberline so that the understory is a mixture of bluegrasses (Poa spp.), sedges (Carex spp.) and various forbs from the alpine zone and other grasses from the lower life zones such as mountain brome, bearded wheatgrass (Agropyron subsecundum), muttongrass, spike trisetum (Trisetum spicatum), Thurber fescue plus naturalized Eurasian species like timothy and orchardgrass locally abundant.

Rocky Mountain National Park, Larimer County, Colorado. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

218. Old-growth stand of Engelmann spruce (subalpine fir is a minor component here)- Understory is dominated by shrubs: blueberry or huckleberry (Vaccinium caespitosum, V. myrtillus= V. oreophilum) with bear-berry or kinnikinic (Arctostaphylos uva-ursi) and current (Ribes spp.) as associates. Hudsonian life zone of C. Hart Merriam.

Rocky Mountain National Park, Larimer County, Colorado. Elevation about 10,000 feet. August. FRES No. 23 (Fir-Spruce Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

219. Finale of a plague- Subalpine Engelmann spruce (dominant)-subalpine fir (associate) forest on west side of Continental divide devestated by outbreak of spruce beetle (Dendroctonus rufipennis ). During the first decade of the Twenty First Century outbreaks of conifer-feeding beetles of various Dendroctonus species occurred across the western half of North America from the west side of the Continental Divide of the Southern Rocky Mountains to British Columbia resulting in deaths of entire populations of Engelmann spruce and subalpine fir as well as lodgepole.

The example shown here was recent death as evidenced by presence of dead needles. Most of the remaining conifers were saplings of Engelmann spruce or subalpine fir (less common were those of lodgepole pine). There were some quaking aspen as well. This was shown to better advantage in the two immediately succeeding slides.

The understorey of this forest tract was shrubby the vast "bulk" of which was dwarfleaf, littleleaf or dwarf huckleberry (see above).

Baker and Veblen (1990) and Veblen et al. (1991) analyzed ecological impacts and plant community changes in Engelmann spruce-subalpine fir due to outbreaks of spruce beetle. Dynamics of spruce-fir subalpine forests were studied and discussed in detail by Aplet et al. (1988). All of these studies were conducted and centered in the Southern rocky Mountains of northcentral Colorado. Simply put, the spruce bark beetle is a native insect that has periodically killed off mass numbers of Engelmann spruce and subalpine fir over extensive regions resulting in elimination of almost all adult trees from spruce-fir subalpine forests. In fact, the Engelmann spruce-subalpine fir forests are greatly reduced over hundreds of thousands of acres for a period of years under these climax forests re-establish through plant succession. Such is the natural dynamics of native vegetation.

Rocky Mountain National Park, Grand County, Colorado. Mid-June (late vernal aspect); approximately 10,000 foot elevation. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

220. Selective grazing by insect and pathogen (or quakies' revenge)- Feeding, mining, and associated activities of high populations of spruce beetle resulted in death of Engelmann spruce and subalpine fir in this subalpine forest. There were a few lodgepole pines in this forest and most of them had been killed by the pine beetle (this was discussed earlier in this chapter). Some of the smaller conifers had survived (so far), but the main tree species surviving were quaking aspen The first of these two slides presented quaking aspen at scale of this forest stand along with a lodgepole pine (far left margin) that had somehow missed pine beetle attack (apparently anyway). The second slide, at smaller spatial scale, featured trunks of quaking aspen along with those of dead Engelmann spruce and also a seedling of Engelmann spruce (center foreground) as one of the first as the next generation of that species. It will be many a year before this little tree grows to adult size of those killed by spruce beetles.

It was described both above and below how forest succession throughout the Southern Rocky Mountains resulted in replacement of quaking aspen by various coniferous species. This species replacement pattern was commonly aided by heavy elk browsing on quakies. In the instance shown here beetles of Dendroctonus species were a natural disturbance that brought about forest retrogression so that once again quaking aspen was set to regain dominance. That is, for a successional period until Engelmann spruce and subalpine fir as the climax dominant and associtae species, respectively, once again reclaimed this forest range site and reigned supreme.

Rocky Mountain National Park, Grand County, Colorado. Mid-June (late vernal aspect); approximately 10,000 foot elevation. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

221. Beetled blessing- Interior of an Engelmann spruce (subalpine fir as associate conifer) subalpine forest with an herbaceous understorey following death of almost every tree by spruce (or spruce bark) beetles. This stand had developed on the upper elevational edge of a wet meadow where drier soils were more favorable for establishment of conifers. The dominant herbaceous species of this understorey was blue wild rye (Elymus glaucus) with bluejoint or Canada reedgrass (Calamagrostis canadensis) as the associate species. There were also some bluegrass species including the naturalized Eurasian Kentucky bluegrass (Poa pratensis) as well as Carex species, especially beaked sedge (C. rostrata). The most common forb was the invasive, naturalized Eurasian weed, common dandelion (Taraxacum officinale), which appeared to be increasing to a critical stage of invasion due to overgrazing by elk. The most abundan native forb was heartleaf arnica (Arnica cordifolia). Rumbaitis del Rio (2006) described a similar herbaceous understorey in an Engelmann spruce-subalpine fir forest following windthrow and salavage logging in the neighboring Routt National Forest.

Impact from beetle-caused death of conifers on herbaceous species remained to be seen. In the short term it seemed likely that the herbaceous species were likely to benefit from increased resources including light, soil moisture, mineral nutreients, etc. These effects was being and will continue to be compounded by heavy elk grazing/broswing. It was undeniable--to this rangeman anyway--that overgrazing by elk had already permitted heavy invasion by the alien common dandelion due to reduced vigor and competitive ability of blue wildrye and bluejoint. Even feces of elk could have some influence on this forest range. Elk dung was widespread throughout this stand and in adjoining stands of subalpine forests (see again above) and wet meadows.

Rocky Mountain National Park, Grand County, Colorado. Mid-June (late vernal aspect); approximaately 10,000 foot elevation. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

222. Subalpine forest- Physiogonomy and structure of a consociation of subalpine fir on east side of Continental Divide at an elevation of roughly 10,000 foot elevation. There was very little of an understorey other than that of subalpine fir regeneration except around the edges where there were some plants of a Salix species, grouseberry or broom huckleberry, and rock ragwort (Senecio fremontii).

Burns and Honkala (1990) explained that the biological range of subalpine fir includes "the coolest and wettest" of forest habitats in North America. Subalpine fir has been interpreted as Tolerant to Very Tolerant depending on associated conifer species. It is more tolerant than Engelmann spruce, interior Douglas-fir, lodgepole pine, and quaking aspen with which it is associated in the Southern Rocky Mountains so that it and Engelmann spruce form co-dominant climax forests. At higher elevations and on cooler environments like north slopes subalpine fir sometimes forms single-species stands (Burns and Honkala, 1990). The forest shown here was an example of such a "pure" stand.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect); roughly 10,000 foot elevation. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

223. Local stand of subalpine fir- Interior of a stand of subalpine fir approaching state of old-growth. The lack of an understorey was obvious although it was still early in the growing season (snow cover was still widespread as shown in the slides preceding and succeding this one). There was very limited regeneration of alpine fir in this stand. So procede to the next photograph...

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect); roughly 10,000 foot elevation. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

224. Proud parents looking down of their progeny- Regeneration of subalpine fir on a road cut just below an old-growth stand that provided seed for this (these) cohort(s). Subalpine fir was found to be a poor seed-producer in the Southern Rocky Mountains with years of good seed crops infrequent. In addition seed viability is generally fair at best (Burns and Honkala, 1990). Furthermore seed germination and seedling establishment is typically best on bare soil (such as roadcuts like the one shown here), but subalpine fir has been found to be less exacting in seedbed requirements than associated spceies being able to germinate and establish seedlings even of thick duff under dense forest canopy (Burns and Honkala, 1990). However seedlings are susceptible to heat injury due to low heat tolerance. Subalpine fir is slow-growing with slow growth rates of roots sometimes being a limiting factor in establishment. This species does reproduce asexually by layering under harsher conditions (eg. timberline), but regeneration by layering is unimportant in mature or closed-canopy forests (Burns and Honkala, 1990).

Sexual reproduction is essential for persistence of this species, but subalpine fir is long-lived such that infrequent reproduction is not a major limiting factor of this Tolerant climax conifer.

Incidentially this was the regeneration scene two days before the summer solstice. Cold-tolerance and, conversely, susceptibility to heat damage of subalpine fir was self-evident.

Some trees of Engelmann spruce were in this stand, but it was essentially a population (= single-species stand) of subalpine fir.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect); roughly 10,000 foot elevation. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

225. Same view the next year- The same population of old-growth subalpine fir and their progeny as shown in the immediately preceding photograph one year later and three weeks later than above. Slow growth rate of subalpine fir was obvious, as was the difference a few weeks advance makes in seasonal progression of the annual growth cycle at this elevation and harsh habitat.

Rocky Mountain National Park, Larimer County, Colorado. Early July (early estival aspect); roughly 10,000 foot elevation. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

226. Invasion by the climax co-dominants- Edge of local subalpine mountain meadow (subalpine park)-- made up of various bluegrass and bromegrass species along with some caric sedges as predominant herbaceous genera--being invaded by Engelmann spruce and subalpine fir from adjacent mature, climax spruce-fir forest. This subalpine meadow had been disturbed by by mechanical snow removal (to clear a nearby highway) as well as disturbance by northern pocket gopher (Thomomys talpoides) resulting in local patches of bare soil and accelerated soil erosion. This denudation of herbaceous vegetation (removal of meadow turf) served as "new land" available for invasion of the climax co-dominant conifers of an adjacent Engelmann spruce-subalpine fir stand.

The actual potential natural vegetation of this local "park" was not known. Or, stated another way, reason(s) for the "clearing" was unclear. Perhaps this local park was the climax vegetation and the woody invasion an example of range retrogression due to zootic disturbances. Alternatively, previous human activity such as building the highway may have caused loss of the spruce-fir forest as the climax vegetation so that more recent disturbance (perhaps coupled with a local seed source) facilitated re-establishment of climax subalpine forest. Either way, subalpine fir and Engelmann spruce were invading the disturbed subalpine meadow.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect); roughly 10,000 foot elevation. Subalpine fir-Engelmann spruce forest was assumed to be the climax vegetation on this range site for purposes of this discussion of forest types in this chapter. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

227. Hole in the canopy- A wind-downed over-ripe (past peak maturity and senescing) Engelmann spruce in an old growth Engelmann spruce-subalpine fir forest resulted in a gap in the otherwise closed crown layer of this range vegetation. With some light reaching the ground surface (understorey) low-growing plant species were able to utilize this precious (limited and essential) resource. Strictly speaking light is one of the abiotic ecological factors that is both a resource and a condition of plant growth. Given the limitedness of light in this forest ecosystem light was most obvious as a resource in development (succession) of this forest community.

The featured recently downed Engelmann spruce was only the latest tree to fall. Note logs of previously fallen adult trees in the second of these two slides (and in the slide of the next caption). It was not known if wind, lightening, or gravity had been the "grim reaper" of earlier downed trees.

Most of the lower community layer was the almost-dwarfish shrub, broom huckleberry or grouse berry (Vaccinum scoparium), with the forb, heartleaf arnica (Arnica cordifolia) the associate species. This lower layer was shown in the next two photographs. Examples of broom huckleberry were presented above.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect); roughly 10,000 foot elevation. Subalpine fir-Engelmann spruce forest was assumed to be the climax vegetation on this range site for purposes of this discussion of forest types in this chapter. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

228. Wind fall- The term "wind fall" was derived from the gathering and use of fallen trees, limbs, branches, etc. for fuel by serf on landed estates during Medieval Times. Villagers and those who lived on estates of the nobility were forbidden from felling of live trees, but people were free to use any dead wood on the ground. Any such fuel so provided by wind or gravity was immediately appropriated by impoverished tenants.

On the forest gap featured in this short section holes in the forest "ceiling" that permitted scarce light to reach the forest floor and rotting logs of fallen Engelmann spruce and subalpine fir that recycled nutrients (fertilized) the soil were a wind fall to whortleberry or broom huckleberry and heartleaf arnica, dominant and associate species of the lowest vegetational layer of this subalpine forest community.

The photographers black shoe in lower left of center provided scale for the low-growing huckleberry and recently emerged heartleaf arnica.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect); roughly 10,000 foot elevation. Subalpine fir-Engelmann spruce forest was assumed to be the climax vegetation on this range site for purposes of this discussion of forest types in this chapter. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

229. Possession of the forest floor, for now- Heartleaf arnica, the locally most abundant species, and broom huckleberry or whortleberry, the overall dominant of this lowest vegetational layer, on the ground surface of a gap formed in an old-growth Engelmann spruce-subalpine fir forest. This was a closer-in view of the forest gap featured and described in the immediately preceding slides and captions.

Although the low-growing shrub and herbaceous composite have the forest floor to themselves at present, the complete coverage of the soil surface by cones of Engelmann spruce (only a few cones of subalpine fir) left no doubt as to which forest citizens would be elected to govern the forest in the very near future. The two climaax dominants of this forest cover type will not have to reach a size of anything near maturity to exclude enough light to result in death of huckleberry and heartleaf arnica seen here and preclude establishment of these two species. In fact, a dense coverage of conifer seedlings will be more effective at excluding light than the lower density and foliar cover of adult trees. (An example of a "dog-hair" stand of subalpine fir seedlings on a roadcut a short distance above this gap was shown above.) Broom huckleberry, heartleaf arnica, and other understorey species will become more plentiful under the mature trees. All of these plant species are components of the climax vegetation.

Heartleaf arnica is a rhizomatous perennial composite that is capable of efficient ground coverage and capture particularily on newly available soil following logging, burning, or natural tree fall as shown here (Hermann (1966, p. 289). One of the most interesting features of Arnica cordifolia is its reproduction solely by apomixis and absence of sexual populations yet with "great variability" resulting from "many slightly different apomictic clones" (Weber, 1990, ps. 72-73). (An example of an adult plant of hertleaf arnica in full bloom was shown below in this chapter.) lBotany and Ecology are such intriguing disciplines. Vegetation is not a dull subject to those blessed with the gifts of awe and wonder.

Rocky Mountain National Park, Larimer County, Colorado. Mid-June (late vernal aspect); roughly 10,000 foot elevation. Subalpine fir-Engelmann spruce forest was assumed to be the climax vegetation on this range site for purposes of this discussion of forest types in this chapter. FRES No. 23 (Fir-Spruce Ecosystem).K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

230. Ecotone between subalpine fir-Englemann spruce forest and limber pine forest (or woodland)- These two photographs were of transitional forest vegetation contiguous with the Englemann spruce-subalpine forest containing considerable cover of limber pine (designated by the author as a "transition" between the two forest cover types or a "combination forest") discussed in the immediatley preceding four slides and the limber pine forest cover type displayed in the following four slides.

Ecotones are transitions or transition zones between two or more different plant communities. There are several forms of ecotones with various kinds of edges or borders. Of course, ecotones are themselves the overlapping borders of adjacent plant communities or vegetation types. Sometimes edges are sharp or discrete. Other forms of ecotones are more duffuse or "blended" over a wider space.

The forest community presented four slides back was one stage or one step in vegetation from Englemann spruce-subalpine fir forest to limber pine woodland. The vegetation seen in these two slides was the next spatial stage in this transition from spruce-fir forest to limber pine forest. This transition was largely a function of elevation and aspect in which limber pine became increasingly more abundant with increasting elevation and progression from a southeastern to a north slope. The forest community now presented was more of a true "mixture" of the two kinds of Rocky Mountain subalpine forest, two kinds of forest range.

It is important that students master the concept of ecotones (vegetational transition zones). Natural vegetation seldom grows in communities as discrete as populations of crops in farm fields or forest plantations.The vegetation displayed here was a good example of a forest ecotone (one in which elevation and slope were determining factors). Ecotones are one of the first examples that shows beginning students that Vegetation Science lacks the accuracy and precision of the physical-chemical sciences and engineering. Vegetation often comes comes in "shades of gray" when resource management demands (or appears to demand) a dichotomous choice.

The first slide showed some young Englemann spruce (eg. the rightmost foreground conifer and several of the spire-topped trees in the background) and common juniper (the small flat plants in the left foreground) intermixed with limber pine. The second slide had a higher proportion of limber pine. These were younger pines with open, widely branching crowns that grew on a south slope. These should be compared (contrasted) to the nearly naked, wind-pruned spires shown below (immediately following the next slide) in a pure limber pine stand on the north slope of this same mountain top. Common juniper was also present (and more conspicuous) in the second slide in the left foreground.

Rocky Mountain National Prk, Larimer County, Colorado. South slope (protected from dessicating, defoliating wind). Elevation about 10,500 feet. June. This ecotonal vegetational did not fit neatly into a simple "black or white" unit, but it was more of the limber pine cover type (SAF 219) whereas the preceding was more of the Englemann spruce-subalpine fir cover type (SAF 206). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

232. Limber pine forest or woodland- This was one of the most unique (and picturesque) forest range types found in the Rocky mountain cordillera. A woody plant-dominated community with erie, pole-like skeletons of ancient, wind-pruned limber pines formed an open upperstorey above a mixed understorey composed of a shrub layer and a pronounced herbaceous layer. This vegetation could be interpreted as either a forest or, given the open canopy of these wind-flagged trees, a woodland. Some authors have referred to this community as the limber pine woodland. It was entitled and described as a forest cover type by the Society of American Foresters (Eyre, 1980, ps. 98-99). The prominent shrubby and herbaceous layers of a browsable/grazable understorey made designation of forest range undeniable. Furthermore, it was permanent forest range.

The range component or feature of this vegetation is not limited to certain seral stages or to climax. Rather, there is a range resource throughout the entire sere (successional sequence) of vegetational development. Limber pine woodland is, however, the climax range vegetation on most forest sites on which it developed and persisted as subalpine forest, especially as krumholtz or elfinwood. Shelford (1963, p. 163) concluded: "Bristlecone or limber pine forms an edaphic climax on dry rocky slopes and windswept ridges".

On this north slope immediately below timberline, wind and wind-driven frozen precipitation (ice, sleet, snow) not only flagged the mature trees but killed back recent foliage on the lower brances of limber pine (eg. lower left foreground). Fallen dead trees so littered the ground that it was possible for man and beast to go from one to another without stepping on the ground. The short cool summers retarded microbial decomposition thereby allowing accumulation of limber pine logs among the slow-growing pines of younger generations and the slowly dying patriarchs of earlier cohorts.

The shrub layer was a combination of prostrate, wind-pruned limber pines as well as numerous kinds of willows and Vaccinium species like broom huckleberry or grouse whortleberry (displayed above). The herbaceous layer consisted of various species commonly found on the alpine elevations above the limber pine zone. Some of these were listed in the next caption and presented individually below.

Wind-polished trunks were on both dead and live limber pines (see succeeding slides also). Compare the flagged, almost krumholtz form of these limber pines growing on a north slope to those on the south side of this same mountain top that were shown in the three photographs immediately preceding the limber pine slide above.

Rocky Mountain National Park, Larimer County, Colorado. North slope. Elevation roughly 11,000 feet. June. FRES No. 26 (a complex of lodgepole, limber, bristlecone, and whitebark pine types). K-8 (Lodgepole Pine-Subalpine Forest). SAF 219 (Limber Pine). Bristlecone Pine-Limber Pine Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

233. Limber pine woodland range type- Physiogonomy, structure, and composition of a subalpine limber pine forest or woodland range cover type with well-developed herbaceous, shrub, and tree layers. Herbaceous species included bronze-scale sedge (Carex chalciolepis), blackroot sedge (C. elynoides), interior bluegrass (Poa interior), blackheaded daisy (Erigeron melanocephalus), and alpine clover (Trifolium dasyphyllum). Dominant shrubs were wind-hedged (one form of krumholtz) limber pines and willows of creeping habit due to wind-pruning and/or genetic morphology.

Two features worthy of note: Number one was that wind-scoured whitish bark was on both live pines as well as persistent in small patches on dead pines. Number two was abundant regeneration of limber pine which indicated that this forest or woodland of an Intolerant species was: 1) climax and 2) uneven-aged. Apparently this north slope habitat had at least two major or general impacts. There was more moisture from winter storms and the generally more mesic soil conditions of a north slope environment. There was also more wind which in winter was cold wind (either dry when "northers" came through or wet winds that drove frozen precipitation with force enough to defoliate and "flag" trees). Various combinations of these factors allowed an open canopy and wide spacing of adult limber pines which enabled germination and development of pine seedlings. The combined affect of wind-dessication and wind-driven ice and snow effectively pruned back crowns of mature pines enough to maintain an open forest floor. This in turn allowed reproduction of limber pine and development of a two- or even three-layered understorey. This understorey was comprised of young limber pines and willow species plus with many grasses, grasslike plants, and forbs common on the the alpine ecosystem above and adjoining this limber pine-dominated climax community.

Rocky Mountain National Park, Larimer County, Colorado. North slope. Elevation about 11,000 feet. June. No specific FRES: closest was FRES No. 26 (a complex of lodgepole, limber, bristlecone, and whitebark pine types). K-8 (Lodgepole Pine-Subalpine Forest). SAF 219 (Limber Pine). Bristlecone Pine-Limber Pine Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

234. Composite view of the limber pine forest range type- Structure and composition of a climax subalpine limber pine forest with the three prominent layers of tree, shrub, and herb distinct. Reproduction of limber pine was obvious. This indicated that crown cover and density of adult pines was low enough to permit regeneration of this Intolerant conifer. The uneven-aged structure (= presence of different crown classes) of the limber pine-- the dominant species-- population strongly suggested that this was climax vegetation. It was discussed in the preceding caption how wind had shaped (ie. pruned) the pine crowns thereby allowing establishment of seedlings of the Intolerant limber pine under the "shade" of their parent trees. Wind-burnished bark and dead wood was present on trunks of both living and dead limber pines. Mesic conditions afforded by the north slope on which this vegetation developed was also a requirement for regeneration of the Intolerant dominant.

Plant cover in the foreground was a composite sample of the shrub layer (= woody understorey) throughout this limber pine dominated-vegetation. The leftmost plant, shrub, in the foreground (to the right of the second boulder) was some species of willow that had a creeping habit due to either wind-sculpting and/or genotype (ie. an example of phenotypic plasticity and/or a combination of morphological plasticity and genetically determined shoot form). Willows (species and number of species was not determined) were the associate species of the woody understorey. The dominant of this shrub layer was wind-hedged limber pine (krumholtz or elfinwood form) such that the dominant of the tree or canopy layer, the overstorey, was also the dominant of the woody understorey (ie. limber pine dominated two of the three layers of this plant community). The krumholtz form of limber pine that dominated the shrub layer was represented by the P. flexilis plant(s) adjacent and immediately to the left of the creeping willow (note dead needles due to dessication by wind and wind-driven frozen precipitation).

It was possible that some of the limber pine component of the shrub layer was actually lower limb and branch portions (krumholtz form) of adult pines. Some adult pines of tree form may have developed elfinwood parts below their single, treelike bole that escaped wind-ice pruning and developed under apical dominance. Determination of such phenomenon would require close investigation, perhaps excavation of stumps and upper root systems. Nonetheless, there was unequivocal evidence of abundant sexual reproduction of limber pine in this stand.

Dominant species of the lowest layer of vascular plants, the herbaceous layer or understorey, was not obvious but was apparently blackroot sedge. Other common herb species were bronze-scale sedge, interior bluegrass (and other bluegrass species that were not blooming and could not be identified), composites like blackheaded daisy, and alpine clover. These herbaceous species were some of those most common on the alpine turfs and fell fields conterminous with the limber pine forest. A representative patch of this often turf-like herbaceous understorey was obvious immediately behind the krumholtz willow and limber pine in the foreground.

It was again remarked this climax plant community was permanent forest range: a grazable/browsable understorey was a component of this vegetation throughout the successional sequence of the sere and was not limited to seral stages prior to climax or canopy closure (whichever came first). Obviously the crown classes did not include the point of canopy closure. There was adequate light for understorey development and persistence throughout the entire spatial array of vegetation development.

Rocky Mountain National Park, Larimer County, Colorado. North slope. About 11,000 foot elevation. June. No specific FRES: closest was FRES No. 26 (a complex of lodgepole, limber, bristlecone, and whitebrk pine). K-8 (Lodgepole Pine-Subalpine Forest). SAF 219 (Limber Pine). Bristlecone Pine-Limber Pine Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

235. Limber pine forest cover type- Climax limber pine forest or woodland (depending on how crown canopy, aerial cover, tree spacing, and density, is interpreted) with a permanent grazable/browsable understorey. Structure and composition of this permanent forest range was evident in this slide. The were three prominent layers: tree, shrubby, and herbaceous. The herbaceous layer consisted of grass, grasslike plant, and forb species abundant of the alpine range ecosystem conterminous with this subalpine limber pine forest. Herb species included blackroot sedge (closest thing to the herbaceous dominant), bronze-scale sedge, bluegrass species, composites, and alpine clover.

The shrub layer was dominated by the wind-sculpted elfinwood form of limber pine with species of willow as associates. Interesting, the same species-- limber pine-- dominated two of the three layers of vascular plants in this vegetation. It was certainly the limber pine cover type.

Death of pine leaves by dessication on this wind-swept north slope was pronounced. Accumulation of dead and down timber of limber pine was due to slow rates of decomposition on this cold site. Even though the north slope aspect created a mesic site much of the annual precipitation comes as snow (or ice) when temperatures are too low for rapid rotting of wood and at this elevation the warm season is too short for much microbial reduction.

Rocky Mountain National Park, Larimer County, Colorado. North slope. Elevation approximately 11,000 feet. June. No specific FRES: closest was FRES No. 26 (a complex of lodgepole, limber, bristlecone, and whitebark pine types). K-8 (Lodgepole Pine-Subalpine Forest). SAF 219 (Limber Pine). Bristlecone Pine-Limber Pine Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

236. Blackroot sedge (Carex elynoides)- This cespitose sedge appeared to be the most common plant in the herbaceous layer of the limber pine woodland presented in the preceding five slides. The carices traditionally have not been accorded coverage like that of the more economically important grasses (or trees), but some outstanding references are available. For C. elynoides refer to Hermann (1970, ps. 204-205) and Hurd et al. (1998, ps. 108-109).

Rocky Mountain National Park, Larimer County, Colorado. North slope in understorey of limber pine woodland; roughly 11,000 foot elevation. June.

237. Bronze-edge sedge (Carex chalciolepis = C. heteroneura var. cahlciolepis )- This is a showy sedge (by sedge standards). It was both abundant and prominent in the understorey of the tract of limber pine woodland discussed above. Standard reference, including forage value, for this Carex species was Hermann (1970, p. 313-314).

Rocky Mountain National Park, Larimer County, Colorado. North slope in understorey of limber pine at about 11,000 feet. June.

239. Blackheaded daisy (Erigerion melanocephalus)- There are several species of Compositae in the alpine ecosystem and adjoining subalpine communities such as the limber pine forest. Blackheaded daisy was the most common forb in the tract of limber pine woodland presented above. Erigerion species are more commonly known as fleabanes. This species was not discussed in Notes on Western Range Forbs (Hermann, 1966), but can be found in standard references including field guides and florae.

Rocky Mountain, Larimer County, Colorado. North slope limber pine climax forest; elevation roughly 11,000 foot. June.

240. Alpine clover (Trifolium dsayphyllum)- There are three, perhaps, four species (depending on personal view) of Trifolium species in the alpine turf and fellfield types and adjoining range types (eg. limber pine forest range) that closely resemble each other. Alpine clover is common, even locally dominant, in all of these high elevation plant communities. The examples shown here grew in the understorey of the limber pine forest tract presented above.

Rocky Mountain Ntional Park, Larimer County, Colorado. North slope forest understorey at approximately 11,000 feet. June.

242 Quaking Aspen (Populus tremuloides) grove— the tree growth form of "quakies" producing an obvious forest type with an understory that is almost exclusively forbs, including poisonous ones. Note the infamous false-hellebore or corn lily (Veratrum californicum) which causes the cyclops lamb tetrogenicity when eaten by ewes on the fourteenth day of gestation. The toxic western larkspur (Delphinum occidentale) is also common , but so are valuable forage forbs like Richardson's geranium (Geranium richardsonii).Routt National Forest, Jackson County, Colorado. July, peak estival aspect.

There are no FRES or Kuchler Numbers for this cover type. FRES No. 19 (Aspen-Birch Ecosystem) would be a rough guide, but all the Kuchler equivalents therein are seral stages whereas quaking aspen does on some sites, as illustrated here, form populations that are not seral. On other sites quakies are clearly seral to spruce and fir climaxes such as those shown in the four preceeding slides. In cases of the latter, quaking aspen can exist as a fire-maintained type. The quaking aspen parkland as representedhere covers vast acreages in parts of Canada and the inland Northwest of the United States where it is scattered throughout or among grassland communities often as a boundary between grassland and the boreal coniferous forest or, at lower latitudes, Douglas-fir and pines like lodgepole and ponderosa. Quakies comprising most of these more northern aspen groves are shrubs. The stature and growth form of those quakies qualifies as a shrubland cover type and could as correctly be shown under the shrublands section. All quaking aspen types were placed here for convenience and to avoid confusion. SRM 411 (Aspen Woodland).SAF 217 (Aspen). Populus tremuloides subclimax Association in ecotone between Douglas Fir-White Fir (Mixed Conifer) Series and Yellow Pine Series of Brown et al. (1998). Southern Rockies- Crystalline Subalpine Forests Ecoregion, 21b (Chapman et al., 2006).

244. Quaking aspen grove- This is another example of a grove of quakies showing the physiogonomy and structure of a relatively young stand. It was growing just above a small stream at the base of a large hillside covered with a mountain big sagebrush (Artemisia tridentata subsp.or variety vaseyana) shrub steppe. Species in the aspen understorey were primarily those of the sagebrush shrub steppe including mountain big sagebrush several individuals of which were growing in the center foreground at the edge of the grove.

Grand County, Colorado. June. It was explained in the first caption of this series that there was no specific FRES or Kuchler Number for quaking aspen. FRES No.19 (Aspen-Birch Ecosystem) was as close as possible. SRM 411 (Aspen Woodland). SAF 217 (Aspen). Populus tremuloides subclimax Association in Douglas Fir-White Fir (Mixed Conifer) Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

245. Interior of quaking aspen grove- Inside the grove of quaking aspen shown in the preceding slide. Species in the herbaceous understorey included timber oatgrass (Danthonia intermedia), Idaho fescue (Festuca idahoensis), cheatgrass (Bromus tectorum), Sandberg bluegrass (Poa secunda= P. sandbergii), and an unidentified species of lupine, all of which were at prebloom stage. Growing immediately adjacent to the boundary of the grove and adjoining mountainbig sagebrush steppe (ie. along the edge of the two range cover types) was squirreltail bottlebrush (Sitanion hystrix) and the prominent forb, false helleborne. The latter was introduced in the first slides of this series.

Grand County, Colorado. June. Approximate FRES No.19 (Aspen-Brich Ecosystem). SRM 411 (Aspen Woodland). SAF 217 (Aspen). Populus tremuloides subclimax Association in Douglas Fir-White Fir (Mixed Conifer) Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

In the author's observations (emperical not experimental) the understorey of quaking aspen groves varies greatly, even over remarkably short distances, depending on the species in adjacent or proximate plant communities where the surrounding vegetation is the dominant cover type of that general location. This observation obtains for scattered groves of quaking aspen in a matrix of other range type(s) and not where aspen is the general dominant range type for that location (not for instance where aspen parkland covers vast expanses and is the regional climax, the climatic climax at regional scale). Where aspen groves occur at elevations at or just above the ponderosa pine zone the aspen understorey tends to have the less mesic grasses and forbs of the pine forest whereas when aspen grows at higher elevations up into the Englemann spruce-subalpine fir forest the aspen grove understorey has the more mesic and shade-tolerant herbaceous species of the latter forest. (Or, as shown immediately below, quaking aspen is seral to the spruce-fir forest.). In the preceding example where the small aspen stand grew in a matrix of mountain big sagebrush steppe the dominant understorey species were those of the shrub steppe.

The successional status of quaking aspen for a given habitat (ie. on a specific sere) may be even more important in determining species composition and structure of the understorey. Aspen groves (or even the larger spatial scale of aspen woodland) might have a different understorey depending on whether aspen is a seral or the terminal stage of plant succession on a given range site or habitat type. An example was shown in the next slide which can be compared with those of other groves in this series.

An excellent source on maintenance of quaking aspen is the symposium proceedings compiled by Shepperd et al. (2001).

246. Stand of quaking aspen being invaded by Englemann spruce and Colorado blue spruce- The successional status of quaking aspen is very much site-dependent, a fact that has often been overlooked when arguments have arisen where aspen was replaced by other species. Yes, indeed sometimes such replacement was due to improper management as when overbrowsing by livestock or big game resulted in loss of quakies and their replacement by herbaceous species or where underburning (or complete fire exclusion) allowed less fire-adapted shrubs and trees to invade stands and ultimately replace the aspen.

Ecologists may never understand completely the dynamics of such vegetational development. It is known that quaking aspen is the potential natural vegetation or climax where it forms extensive parklands in more northern regions. At the other end of plant succession aspen is sometimes a pioneer species that colonizes disturbed areas. This latter pehnomenon would apply to secondary not primary succession. It is also known that on some habitats quakies are seral to coniferous forests. One of the best known examples of this latter successional pattern is where (again, on some sites) quaking aspen is seral to the climax Englemann spruce-subalpine fir forest. That condition was shown in this photograph.

An old stand of quaking aspen was being invaded mostly by Englemann spruce and Colorado blue spruce with some lodgepole pine (Pinus contorta var. latifolia) and even a few limber pine (Pinus flexilis) also appearing to become established. (The pine saplings were too small to be relicts of the same age as the aspen.) Quaking aspen (along with cottonwoods) was ranked by the Society of American Foresters as Very Intolerant, the least tolerant of any western hardwood, whereas Englemann spruce was ranked as Tolerant and subalpine fir was Very Tolerant (Wenger, 1984, ps. 3-4). Where the site potential will support spruce-fir forest it is inevitable that these species will eventually replace the quakies, unless of course repeated disturbances like fire and avalanches prevent progression of plant succession.

This forest vegetation was on an east slope and there was a spring or hillside seepage at the lower edge of the forest where it was advancing downslope and invading a wet meadow. This forest site, especially the wettest microsite, was extremely favorable to blue spruce.

The two young conifers in the far right foreground were Englemann spruce and the conifers in the background were the famed Colorado blue spruce. No subalpine fir were found and there were no adult subalpine fir trees present within the immediate vicinity. Blue spruce were more common downslope and Englemann spruce more abundant upslope from this forest stand, as was the limber pine which grew in association with the spruce-fir forest at higher elevations as the spruce and fir gave way to limber forests.

Readers can find a straightforward discussion of the natural history of quaking aspen groves in Mutel and Emerick (1992, ps. 141-151). These authors presented a very readable account of this community (both plant and animal) that will help students understand basic structure, composition, and dynamics of this forest or shrubland type.

Rocky Mountain National Park, Larimer County, Colorado. Approximate FRES No.19 (Aspen-Brich Ecosystem). SRM 411 (Aspen Woodland). SAF 217 (Aspen). Populus tremuloides subclimax Association in Douglas Fir-White Fir (Mixed Conifer) Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

247 Some of us grew; some of us only grew older- The same photgraphic point of a grove of quaking aspen being invaded by Engelmann and Colorado blue spruce as shown above-- sixteen years earlier. Oviously the conifers have grown (some of them considerably) while the existing aspen shoots have senesced (slowily dying following shoot maturity) and only a few new aspen shoots were produced (ie. senescence combined with lack of regeneration). This asexual reproduction of the "quakies" was limited to the perimeter of the grove so that there was some incremental expansion of aspen around outer edges while conifers were "banishing" (overwhelming) the senescing aspen shoots. With on-going plant succession conifers (primarily Colorado blue spruce) were in the process of "ousting" the quakies" and laying claim to the sere, specifically the interior of the aspen grove. Even previously existing lodgepole pine had grown and new seedlings and saplings of lodgepole pine had invaded (established).

Growth rate of all these trees had been relatively slow in the continental climate of the Southern Rocky Mountains with long, cold winters and short, cool summers. This slow growth of all tree species can be easily ascertained by comparison of this photograph with the one of sixteen years earlier seen immediately above. Nonetheless, increase in size of trees (height, diameter breast height, canopy cover) was visibly greater in conifers, especially blue spruce, than in quaking aspen. Furthermore (and ultimately more important) was the greater rate of regeneration of conifers relative to aspen, the latter of which was nearly nil. It was obvious what the successional outcome would be over time, the time period of forest development. That is, of course, unless some other factor(s) changed the trajectory of forest community change (see below).

Quaking aspen, as was detailed above, is a clonal organism. This aspen grove was mostly--if not exclusively-- one tree (one clone= one genetic individual, one genotype) of numerous shoots. The "trunk" of the tree is a rhizomatous system with "trees" in the grove being shoots ("limbs" or "branches") off of the underground "trunk". In absence of a regenerating "disturbance" such as fire or avalanche the aging aspen clone became a maturing (and slowly dying) grove of old "trunks" (shoots).

Meanwhile, individuals of the two coniferous species (single trees of a genotype, a unique genetic individual conifer) continued to grow and increase in size and crown cover (trees got taller with larger crowns of bigger and more branches with more needles), plus with some sexual reproduction (seed production). In the sixteen year interim between time of forest vegetation shown in the preceding photograaph and that seen in this (and subsequent) photographs Colorado blue spruce had "outdistanced" Engelmann spruce and had come to dominant the developing forest. More lodgepole pines had established (invaded) in the perimeter of the advancing aspen grove.

As new shoots of "quakies" grow and mature around the perimeter of the grove, Colorado blue spruce and Engelmann spruce along with loegepole pine will invade that land and displace (replace) aspen through plant succession.That is, this successional sequence, this profression of plant succession and conifer forest development, would continue if some other factor or phenomenon did not interfere. More on that phenomenon below ...

Rocky Mountain National Park, Larimer County, Colorado. Early July (earrly estival aspect). Formerly this forest vegetation fit the following approximate units: FRES No.19 (Aspen-Brich Ecosystem), SRM 411 (Aspen Woodland), SAF 217 (Aspen),. Populus tremuloides subclimax Association in Douglas Fir-White Fir (Mixed Conifer) Series of Brown et al. (1998). Sixteen years later the forest had undergone progression (plant succession) on the sere to the extent that the range vegetation now more closely fit the following units: FRES No. 23 (Fir-Spruce Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 216 (Blue Spruce) locally at elevation just below the zone of SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

248. Coming on strong and slowing going out- Exterior (general) and interior views--first and second photographs, respectively--of a grove of quaking aspen, the adult shoots of which were senescing faster than reproduction (which would have mostly asexual) of new shoots necessary to replace mature ones and maintain the population. Concurrently, Colorado blue spruce was on tract to become dominant (with some Engelmann spruce and lodgepole pine as associate species) as existing trees of these species advanced in age, size and relative canopy cover as well as having sexual regeneration so as to expand their presence and cover into parts of the grove previously unoccupied by them.

This upper montane, east-slope forest was moving via plant succession from an aspen grove to a conifer forest dominated by blue spruce. There had been some regeneration of aspen--as seen in foreground of first photograph--on the outer margin of the grove, but there had been essentially no (certainly very litttle net) regeneration of aspen in the interior of the grove. Meanwhile, the conifers (again, mostly Colorado blue spruce) were continuing to grown, reproduce, and increase their relative canopy cover throughout this forest community including at the margin or outer extremity of the forest as seen clearly in the second photograph. Conifers at the exterior of this forest (second photograph) where it was advancing into (invading) a wet meadow ranged from larger seedlings to saplings and included both blue spruce and lodgepole pine. New aspens (clonal shoots or sexual seedlings/saplings) were absent from the leading edge. The "quakies" have had it--unless something changes.

Rocky Mountain National Park, Larimer County, Colorado. Early July (earrly estival aspect). FRES No. 23 (Fir-Spruce Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 216 (Blue Spruce) locally at elevation just below the zone of SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

249. Overwhelmed- Adult Colorado blue spruce grrew up through quaking aspen which senesced and died out (perhaps partially reduced due to competition with the conifers) so that the previus aspen grove (SAF cover type 217 SRM cover type 411) was converted through plant succession to blue spruce fores (SAF cover type 216).

This was a local stand in the larger aspen grove shown above. It was the part of tht grove that had moved farthest on this sere (forest site) toward coniferous forest and away from the previous advanced seral stage of the quaking aspen cover type. A spring or water seepage characterized and distinguished this local habitat of the sere (most precisely it was a wet hill slope microsite) so that it was the wettest part of the (former) aspen grove.

Eyre (1980, ps. 95-96) interpreted the blue spruce forest cover type as "subclimax" that typically was replaced (succeeded) by the Engelmann spruce-subalpine fir cover type.It was further explained that the blue spruce type (SAF 216) was a relatively restricted type that was present as climax only along streams, where forest edges met meadows, on "low moist soils", or "subirrigated slopes", The latter three of these conditions existed on the land on which this stand of maturing blue spruce had developed. Furthermore, this forest vegetation was on an east slope so as to be more mesic to begin with.

It was predictable that successional advance from quaking aspen to blue spruce-dominated coniferous forest would be most rapid on this wet hillside. Wet edaphic conditions probably also explained why blue spruce was dominant instead of Engelmann spruce (or even subalpine fir) which dominated higher elevation forests immediately above this developing forest.

Rocky Mountain National Park, Larimer County, Colorado. Early July (earrly estival aspect). FRES No. 23 (Fir-Spruce Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 216 (Blue Spruce) locally at elevation just below the zone of SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

250. Come and go, give and take- Another view of the advancing margin of a mesic, high montane forest on which conifers (namely Colorado blue and Engelmann spruce with some lodgepole pine) were replacing quaking aspen with the progression of plant succession (ie. aspen is seral to a blue spruce climax or subclimax on this forest site). This forest was on an east slope and invading a wet meadow.

Immediately obvious was the absence of any quaking aspen reproduction (neither asexual or clonal shoots nor new trees as seedlings or saplings) accompanied by abundant reproduction of both Colorado blue and lodgepole pine (foreground). Higher up on this east slope Engelmann spruce of sapling and small pole sizes were abundant.

Engelmann spruce and lodgepole pine were also in trouble. In spite of regeneration of these two conifers a widespread catastrophic outbreak of spruce (or spruce bark) and pine (pine bark) beetles (Dendroctonus species) had decimated populations of Engelmann spruce and lodgepole pine throughout this region. Plus, bark beetles were continuing to kill many and, in local areas, most of the few remaining trees. Some beetle-killed, pole-size Engelmann spruce were visible in this photograph (midground). These Dendroctonus species are native insects. For millenia bark beetles have functioned as natural disturbances that displaced climax conifers and set secondary plant succession into motion. Impact of the Dendroctonus decimation on Engelmann spruce-subalpine fir, lodgepole pine, and ponderosa pine climax forests in this area were described earlier in this chapter.

Bark beetle herbivory--again, a natural disturbance--was a phenomenon that was favorable to the seral quaking aspen. Examples of this were shown above. Aspen were still "not out of the woods", however, because excessive defoliation, overbrowsing by elk had been a major factor in preventing reproduction (sexual and asexual) of quaking aspen along the advancing edge of this forest. The result was that woody invasion of the wet meadow downslope from the established forest had been limited to that by conifers. The natural pattern of forest succession was interrupted by elk overbrowsing so that blue spruce, Engelmann spruce, and lodgepole pine were establishing without having to replace aspen (and perhaps without facilitation by this seral species).

Another sure sign of heavy browsing by elk were the many glaring scars on aspen trunks resulting from "barking" (bark feeding) by this generally grass-preferring cervid. This showed up well on this photograph. Close-up photographs of bark scars were presented in the immediately two succeeding slides.

Herbivory by bark beetles had slowed the advance (and were still in process of slowing) invasion of the wet meadow except Colorado blue spruce had yet to be impacted by Dendroctonus species. Smaller trees (seedlings and saplings) of affected conifer species typically are not fed on by bark beetles. Hence, conifers of these age/size classes were still alive at advancing (invading) edge of this forest (foregroung of photograph). Would bark beetle populations subside to normal level so that these young trees would reach adulthood? Would the wet meadow ecosystem survive or would it become a coniferous forest (or maybe woodland or savanna)? Only God knows and time alone will tell.

Rocky Mountain National Park, Larimer County, Colorado. Early July (earrly estival aspect). FRES No. 23 (Fir-Spruce Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 216 (Blue Spruce) locally at elevation just below the zone of SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

251. Elk-carved totem pole- Bark scars on trunks of adult quaking aspen at the edge of a senescing aspen grove that had expanded down an east slope and invaded a wet meadow. Elk prefer herbaceous plants, especially grasses and grasslike plants, but they are opportunistic and diverse feeders that readily consume browse. Quaking aspen are a preferred elk browse plant. Leaves, buds, and young shoots appear to be the more preferred parts of aspen, but elk readily feed on aspen bark. Bark-feeding has appeared to be more widespread under conditions of high stocking rates of elk, especially at levels regarded as overbrowsing.

Feeding on new clonal shoots by elk had unquestionably retarded--in fact, at this location elk overbrowsing had completely precluded--aspen regeneration. In combination with ample reproduction of blue spruce, lodgepole pine, and, at lower rates, Engelmann spruce this grove of quaking aspen was moving rapidly via plant succession to a blue spruce forest. (This was explained immediately above.)

The small shrubs in these two photographs (especially the first slide) were alder (Alnus incana), a riparian species that was locally abundant in the wet meadow at the edge of and immediately below this aspen grove that was becoming a forest dominated by Colorado blue spruce. The yellow-flowered forb on ground layer was heartleaf arnica.

Commonly, quaking aspen quickly replaces bark removed by browsing elk. That was evident from these two photographs. Extremely "deep browsing" by elk below bark and into cambium results in much slower healing wounds. Two such wounds were visible in the shoot shown in the first of these slides. Also noteworthy in this slide was the small snag of a sprout (offshoot, clonal shoot) produced by this adult, parent shoot and promptly killed by elk overbrowsing. Again, an overpopulation of elk and consequent overbrowsing had virtually stopped aspen regeneration.

The disgrace of this unacceptable management was on a national park! Rocky Mountain National Park should be opened to sport hunting or else culling by government employees such as personnel of Animal Damage Control in the United States Department of Agriculture. Establishment of native predators, especially the grey wolf (Canis lupus), might eventually result in reduction of elk numbers in Rocky Mountain National Park. If and until such natural population control occurs human intervention by hunting (culling) is absolutely essential for proper use of range resource and the general Southern Rocky Mountain ecosystem.

Degradation of quaking aspen range by elk-overbrowsing has received--quite justifiably--much attention, by range and wildlife researchers. Range investigators found that heavy utilization and, especially, overbrowsing of quaking aspen is compounded by (and conclusions confounded with) invasion of aspen groves by conifers. One thorough study of this dual threat to "quakies" in the Southern Rocky Mountains was conducted in Rocky Mountain National Park and Roosevelt and Arapahoe National Forests, two large neighboring parcels of public elk range (Kaye et al., 2005). In addition to its original research (which demonstrated what any observant and impartial observer already knew), this paper by Kaye et al. (2005) provided a thorough bibliography of this on-going threat to and, on all-too many ranges, loss of natural vegetation and wildlife habitat.

Rocky Mountain National Park, Larimer County, Colorado. Early July (earrly estival aspect). FRES No. 23 (Fir-Spruce Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 216 (Blue Spruce) locally at elevation just below the zone of SAF 206 (Englemann Spruce-Subalpine Fir). Englemann Spruce-Alpine Fir Series of Brown et al. (1998). Southern Rockies- Sedimentary Subalpine Forests Ecoregion, 21e (Chapman et al., 2006).

255. Rocky Mountain locoweed, white loco, or silky crazyweed (or other combinations of these descriptive words)- Oxytropis sericea is another showy forb found in aspen groves and, in particular, along edges of groves. This robust example was growing where an aspen grove adjoined a subalpine mountain meadow. The pigmented spot on the keel of this papilinaceous legume allowed positive identification to this rather varied species.

Oxytropis is one of the more intriguing genera of poisonous range plants, and O. sericea has figured prominently in the saga. Some Oxytropis species and some of the Astragalus species are classic locoweeds, those that induce locoism or loco poisoning. This condition has been the subject of scientific study for over a century. The poisonous principle was identified as a group of indolizidine alkaloids. Summary coverage was provided by Cheeke and Shull (1985, ps.142-146). Recently O. sericea has been implicated in big brisket, high mountain disease, or congestive right hear failure. For detailed recent treatment see Burrows and Tyrl (2001, ps. 594-599). And never forget Kingsbury (1964, ps. 306-311). Investigations with Oxytropis, including O. sericea, have been reported consistently in the peer-reviewed proceedings of the International Symposium on Poisonous Plants. Readers with a joint enjoyment of Chemistry, Botany, and Range Management can get a full-course feast there. The subject of poisonous range plants has long fascinated those who study and report on "things range". The current writer was no exception.

Rocky Mountain National Park, Larimer County, Colorado. June.

256. Inflorescence of elephant head or little red elephant (Pedicularis groenlandica= Elephantella groenlandica)- This is another showy member of the aspen community (and adjoining forest cover types such as interior Douglas fir). Elephant head was described in both Notes on Western Range Forbs (Hermann, 1966, ps. 264-266) and Range Plant Handbook (Forest Service, 1941, p. W143) where it was described as usually no higher than Fair forage for small ruminants. It is very distinctive and adds species diversity to range plant communities. Elephant head is an easy one to learn (and to impress others with one's knowledge of native plants). This one grew along the edge of an aspen grove in association with cowparsnip (Heracleum lanatum).

Rocky Mountain National Park, Grand County, Colorado. Early August.

260. False helleborne or cornhusk lily (Veratrum tenuipetalum)- According to some of the more recent treatments (eg. Weber and Wittmann, 2001) this is the only species of Veratrum found in the Southern Rocky Mountains in Colorado, but previous work (eg. Dayton, 1960) reported that western false hellebore (V. californicum) occurred in Colorado and, in fact, in all of the 11 Western States. From the current author's observation V. californicum has broadly ovate leaves that are clearly distinct from the lanceolate leaves of the individuals shown here which were therefore designated as V. tenuipetalum. More importantly from this authors perspective was presence of the numerous conspicuous lanceolate leaves on the flower stalk of these plants in contrast to the leafless rachis of the ovate-leaved plants displayed immediately below.

The plants presented in this slide were representative of many growing prolificly at the edge of a the last grove of quaking aspen shown above (Grand County, Colorado). The toxicological property of Veratrum species and the resultant phenomenon of the cyclops lamb was noted in the first caption of this quaking aspen series. The most recent comprehensive reference tretment was that of Burrows and Tyrl (2001, ps. 782-787). These authors did not list V. tenuipetalum but discussed all Veratrum species as toxic. They too showed V. californicum as native to Colorado, western Colorado.

Grand County, Colorado, Colorado. June.

261. Western false helleborne or corn lily (V. californicum)- The broadly ovate leaves of these plants appeared to to this author to be distinctly different from those examples shown immediately above. Even more definitive was the presence of the lanceolate leaves all along the flowering stalk of the plants your author labeled V. tenuipetalum immediately above and the leafless rachis of the plants shown here and labeled V. californicum. Yes, indeed these species-- as interpreted by your author-- both grew in northern Colorado and in adjoining large counties, but the two were on opposite sides of the Continental Divide. The species shown here was on the West Slope and the one shown above on the East Slope. Numerous plant species have been distinguished and separated out on the basis of morphological features much less distinctive and obvious that these, and with less geographic separation than that of the Great Divide. In the first comprehensive maunal of Colorado plant life, the classic Flora of Colorado, Rydberg (1906, p. 76) showed both of these species (V. speciosum was used as the synonym for V. Californicum). Rydberg separated them on oval versus oblong-lanceolate petals. This might be a good research project for some budding plant taxonomist.

One last lesson from this example, the author bought a copy of Rydberg's Flora of Colorado that had been discarded from, first, the University of Montana and, next, Montana State University (discarded stamps from both institutions). Do these two universities, both of which have programs in Forestry and Range Management, have another copy? Just because a book is a century old does not mean it is no longer valid or of use. The currently commonplace practice of discarding texts and references by university libraries based on nothing but copyright dates and how many times the book was checked out is as stupid and short-sighted as it is sinful to knowledge. Oh yes, of course there is inter-library loan. This author can tell you about that too.

The specimens presented here grew inside of and all around the quaking aspen grove shown in the first two slides of this series (Routt National Forest, Jackson County, Colorado, July).

Note: Changes in vegetation due to cycles of changing climate was a popular concept among pioneer plant ecologists (eg. sunspot activity was a “pet” theory) a century before some contemporary climatologists and ecologists starting grandstanding and crowing about human-induced “global warming” and possible consequent modifications of vegetation.

267. Edge of the P-J woodland and adjoining grassland seen in two preceding shots- Rocky Mountain maple is conspicuous. In addition to forbs, herbs include the dominant grasses blue grama, bluebunch and western wheatgrassses, and galleta.

Cache National Forest, Cache County, Utah. July. FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Pine Woodland). SRM 412 (Juniper-Pinyon Woodland); SAF 239 (Pinyon-Juniper). Pinyon-Juniper Series in Great Basin Conifer Woodland biotic community of Brown et al. (1998). Wasatch and Unita Mountains-Semiarid Foothills Ecoregion, 19f (Woods et al., 2001).

268. Petered up- Highest elevational limit of pinyon pine-juniper woodland in the Sagre de Cristo range just above the San Luis Valley. Above this foothill elevation (approximately 7200-7300 feet) and on a xeric west slope was the beginning of a rather scant Douglas-fir forest in the Lower Montane zone. Some smaller Douglas-fir trees were growing on the upper slopes and crest of hills seen here. Most of the low-growing shrub were was current. There were also some plants of mountain ninebrk (Physocarpus monogynus) and willow of undetermined species (Salix sp.) lower down by a small stream.

Herbaceous species were lacking suggesting that this hillside range had been overgrazed and severly depleted.

Saguache County, Colorado. Late June.

One of the most biologically diverse (and beautiful) forms of the pinyon pine-juniper woodland is the oak-pinyon pine-juniper woodland. There is great species diversity in the tree, shrub, and herbaceous layers of this southwestern North American range woodland. One of the best geographic locations for this range vegetation is in the Big Bend area of west Texas and northern Chihuhua where floristic elements from the Rio Grande Plains, Edwards Plateau, Southern Great Plains, and Chihuhuan and Trans-Pecos Basin and Range "blend" to produce an amazingly varied and colorful array of range plants in turn responsible for a remarkable development of this range cover type. The mixed oak-Mexican pinyon pine-juniper woodland begins in mountain foothills above basins and bajadas of Chihuhuan Desert and lower foothills of various zonal grassland communities.

There are various range cover types of Chihuhuan Desert and semidesert (Chihuhuan) grasslands below (ie. "down the mountain and out on the flats") the oak-Mexican pinyon pine-juniper montane woodland. These lower, more xeric range types were covered in this publication under Chihuhuan Desert (Shrublands) and semidesert grasslands (Grasslands).

Arguably one of the most accessable examples of the oak-pinyon pine-juniper woodland range type is in the Chisos Mountains. The following set of photographs was taken from Green Gulch in the Chisos Range. Enjoy the scenery.

The first two photographs in this set were landscape-scale views that displayed the physiography of the Chisos Mountains and the physiogonomy, structure, and general composition of their range vegetation.

270. Oak-pinyon pine-juniper montane woodland in the Basin and Range- Deep in the core of the Chisos Mountains a tremendously varied montane range plant community had developed consisting of several layers of vegetation: 1) tree, 2) shrubs that typically formed two layers, and 3) herbaceous plants consisting of tall-, mid-, and shortgrass species as well as forbs. Even the tree layer of this woodland range was more complex than might be expected due to presence of mistletoe as a parasitic epiphyte.

Coniferous tree species included Mexican pinyon pine (Pinus cembroides), the ultimate climax conifer, alligator juniper (Juniperus deppeana), redberry or Pinchot juniper (J. pinchotii), rose-fruit juniper (J. erythrocarpa), and, less commonly, weeping or drooping juniper (J. flaccida). Woody angiosperm species which included some individuals of tree shape and size were mostly Emory oak (Quercus emoryi) and gray oak (Q. grisea). These were common trees within the photographic view shown here. Other oaks in the Chisos that attain tree deminsions and shape yet which were not observed within field of this photograph included Chisos red or Graves oak (Q. gravesii= Q. chesosensis), chinkapin oak (Q. muehlenbergii), and netleaf oak (Q. rugosa). Uncommon to rare species of oaks that have tree form in the Chisos ncluded Chisos oak (Q. graciliformis) and lateleaf oak (Q. tardifolia). Common scrub oak species scattered throughout the range vegetation presented here included gray oak and netleaf oak (two of several oak species having individual plants that grow in either tree or shrub form) and Vasey shin oak (Q. vaseyana= Qpungens var. vaseyana= Q. undulata var. vaseyana). An uncommon but extremely conspicuous tree species was Texas madrone or naked Indian or lady's leg (Arbutus texana) of the Ericaceae, heath or heather family.

Other angiosperm species on this woodland range that usually have shrub shape and size but sometimes attain tree form or size present included three sumac species: lanceleaf sumac (Rhus lanceolata= R. copallina var. lanceolata), evergreen or tobacco sumac (Rhus virens subsp. virens), and littleleaf sumac (Rhus microphylla). One of the most common angiosperm shrubs was golden or yellow currant (Ribes aureum). The most conspicuous shrubs in many spots were the succulent or succulent-like species including various pricklypear cactii (mostly Opuntia englemannii), smooth sotol (Dasylirion leiphyllum), and lechuguilla (Agave lechuguilla).

Grasses were all native perennial species with Eurasian cool-season annuals like cheatgrass (Bromus tectorum) present at trace amounts. Little bluestem (Andropogon scoparius= Schizachyrium scoparium) and cane bluesteem (A. barbinodis= Bothriochloa barbinodis) were grass species of largest mature size. Little bluestem was interpreted as a tallgrass species. Sideoats grama (Bouteloua curtipendula) was the dominant mid-grass species. Chino grama or chinograss (B. ramosa) was the most abundant grass, but cumulative foliar cover of this species was locally less than that of larger-growing species including cane bluestem and sideoats grama. Chino grama was viewed as a shortgrass species of which grass group it was dominant. Other Bouteloua species included blue grama (B. gracilis), hairy grama (B. hirsuta), and, very infrequently, red grama (B. trifida). Threeawn species were also common (and always the taxonomic nightmare) with red threeawn (Aristida longistea= A. purpurea var. longiseta) the major member of this taxon. Mountain muhly (Muhlenbergia montana) was found rarely (and always in battered condition). Overall there was no dominant grass species.

Forbs were inconsequential at the early summer season shown in these photographs.

Green Gulch, Big Bend National Park, Brewster County, Texas. June. FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak varaint of SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

271. Climax vegetation of an oak-pinyon pine-juniper woodland- Foothills of Chisos Mountains and an almost unbelieveable array of range plants in another panaramic view of Chihuhuan or Madrean montane woodland.. The two tallest trees in midground were Mexican pinyon pine. This species has usually been interpreted as the potential natural dominant gymnosperm of this range cover type. Associate coniferous species are the various Juniperus species listed above. The two junipers in the foreground of this slide were redberry or Pinchot juniper. Shrub in right foreground (right corner of photograph) was gray oak. Smooth sotol was also present. Grasses included sideoats and chino grama and cane bluestem.

Green Gulch, Big Bend National Park, Brewster County, Texas. June, early estival aspect (prior to onset of main summer rains and most grass "green-up"). FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak variant of SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al, 1998). Chihuhuan Deserts- Chihuhuan Montane Woodland Ecoregion 24d (Griffith et al., 2004).

272. Inside Chisos Mountain montane woodland range- Details of oak-pinyon pine-juniper woodland vegetation inside a mountain range in the Trans-Pecos Basin and Range. In this "photo-plot" there was a stand of Mexican pinyon pine with a predominantly shrub understorey sotol, pricklypear, gray oak, and golden currant. There were some Pinchot or red berry junipers (eg. needles of one in right foreground). Chino and sideoats gramas were main grasses.

Green Gulch, Big Bend National Park. June, early estival aspect. FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak variant of SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodland Ecoregion 24d (Griffith et al., 2004).

273. Chisos Mountains montane range- A detailed view inside the oak-Mexican pinyon pine-juniper montane woodland. Taller conifers (eg. three in midground) were Mexican pinyon pine. Shrub in right foreground was a much-brancned redberry or Pincot junipe. According to Powell (1988ps. 38-41) redberry and rose-fruit juniper integrade to the extent that there may be a J. pinchotti X J. erythrocarpa hybrid in the Chisos Mountain Basin. Sotol, including a prominently blooming specimen, was a dominant shrub in mid foreground. Pricklypear was another prominent shrub. Major grasses visible were little and cane bluestems.

Green Gulch, Big Bend National Park, June, early estival aspect. FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak variant of SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

274 Chisos Mountains montane woodland range at its pinnacle- Textbook view of vegetation of mixed oak-Mexican pinyon pine-mixed juniper woodland. Three coniferous trees in center (large mature one flanked by two small ones) was Mexican pinyon pine. These in turn flanked by either redberry juniper or rose-fruit juniper (these two species are very difficult to tell apart). Grass species in foreground were cane and little bluestem, sideoats grama, and red threeawn. Sotol also visible, including flower stalk of one in left foreground.

Green Gulch, Big Bend National Park, June, early estival aspect. FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak variant of SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

275. Cool sweep of Chisos Mountains vegetation- First-timers to these hills find it "hard to imagine" that the Chihuan Desert lies less than 2000 feet below these densely wooded Chisos Mountains (and with an array of distinctive zones of grassland vegetation in between), and that such differences in range vegetation are largely a function of elevation and other abiotic factors associated with elevational gradations.

Range vegetation shown here was a mixed oak-Mexican pinyon pine-juniper woodland. This was a climax plant community with a prominent physiogonomy, rich species composition, and complex structure. Major grass species in this "photo-transect" included cane bluestem, little bluestem, sideoats grama, chino grama, and red threeawn. The conspicuous shrubs in foreground were gray oak of characteristic coloration. Larger trees that were visible in the far foreground were Mexican pinyon pine, alligator juniper, Emory oak, and Texas madrone.

Green Gulch, Big Bend National Park, June, early estival aspect. FRES No. 35 ((Pinyon-Juniper Ecosystem). K-21 ((Juniper-Pinyon Woodland). Oak variant of SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

276. Montane browse above the Chihuhuan Desert- Another landscape-scale photograph of the Chisos Mountains with mixed oak-Mexican pinyon pine-juniper montane woodland. This panoramic view featured details of the browse layer of this range cover type. Shrub species in the foreground included (roughly left to right allowing for intermixing of branches) golden currant, smooth sotol, littleleaf sumac, evergreen sumac, gray oak, and Emory oak. The large tree was Emory oak. Primarily redberry or Pinchot juniper, rose-fruit juniper, and gray oak in basin of background.

Big Bend National Park, Brewster County, Texas. June, early estival aspect. FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak variant of SRM 504 (Juniper-Pinyon Pine Wooodland). SAF 239 (Pinyon - Juniper).Oak-Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

277. Rangeman's picture postcard of Chisos Mountain range- At the base of a mountain slope topped with Casa Grande was an all-in-one shot of the oak-Mexican pinyon pine-juniper woodland range type followed by a second slide showing overall species compostion and structure of this montane vegetation. Tallest tree at far left of both photographs was Mexican pinyon pine. Immediately to the right of the pine was an Emory oak with gary oak to its right. Alligator juniper, redberry juniper, rose-fruit juniper, and Texas madrone were the other major trees higher up on the mountain side. In the foreground the understorey was mostly herbaceous with red threeawn the dominant species. Mule deer (Odocoileus hemionus) were nosing around in this rank-growing grass, perhaps looking for acorns or pine 'nuts".

Big Bend National Park, Brewster County, Texas. June, early estival aspect. FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak variant SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series 123.32 of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

278. The dry side and after a fire- "Just around the corner" from the location of the two immediately preceding slides the range vegetation took on this drastic difference in physiogonomy and species composition. Obviously a fire had modified this vegetation in the not-too-distant past. While the fire "opened up" this range plant community it was probably the xeric, predominately west slope that was responsible for a grass-scrub savanna variant form of the oak-Mexican pinyon pine-juniper woodland type.

The centerpiece showy succulent was Havard agave (Agave havardiana) which was, as if on cue, at full-bloom, and already starting to die as it was fulfilling its once-in-a-lifetime role to reproduce sexually therby giving its species another opportunity for natural selection. Red-fruited shrub in right foreground was littleleaf sumac. Top-killed and resprouting tree at far left foreground was Emory oak. Shrub to left front of agave was redberry juniper which had also been top-killed and subsequently resprouted. Redberry or Pinchot juniper is one of the very few Juniperus species to resprout, a rare characteristic of conifers. The largest tree (directly behind agave) on the slope was a drooping or weeping juniper. This plant was not bearing cones, but even from a distance the descending, dead, outer branches (undoubedly fire-killed) gave this species away as far as it could be seen. This Juniperus species does not resprout, but this particular tree had attained size enough to withstand-- minus portions of some outer branches-- the heat of what was likely an upslope, hot, heading fire.

Predominate grasses were little bluestem, cane bluestem, and sideoats grama.

Casa Grande crowned the spectacle.

Chisos Basin, Big Bend National Park. June. FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper -Pinyon Woodland). Oak variant of SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak -Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004)..

279. Chisos Basin range- Range vegetation in Chisos Basin. The rich flora of climax oak-Mexican pinyon pine-juniper woodland was on display in this view across the Chisos Basin of the Chisos Mountains. Center conifer was Mexican pinyon, the climax dominant gymnosperm. Two Havard agaves in full-bloom "accompanied" the pine. The shrub in center foreground was of the Emory oak scrub form. Grasses included sideoats grama, cane bluestem, red threeawn, and chino grama.

Chisos Basin, Big Bend National Park, Brewster County, Texas June, early estival aspect. ,FRES No. 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak variant of SRN 504 (Juniper- Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series, 123.32, of Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24d (Griffith et al., 2004).

280. Range vegetation of Chisos Basin- Detail of range plants in Chisos Basin. A mixture of herbaceous and woody range plants were easily identified in this photograph. Englemann pricklypear and smooth sotol were in left and right foreground. Sideoats grama, a decreaser and the climax dominant on this location, was in center foreground. Other major grass species included cane bluestem, chino grama, and red threeawn. Shrubs of gray oak were in center midground. Foremost conifer (rounded crownin right midground was redberry or Pinchot juniper. Conifer behind and to left of juniper was a young Mexican pinyon pine. and shrubs.

There were not many forbs growing at the early summer season at time of this photograph. As to be expected most forbs were composites. Most of the dead shoots of forbs were cool-season annuals of the Umbelliferaeh

Chisos Basin, Big Bend Natioal Park, Brewswter County, Tesas. June, early estival aspect. FRES No., 35 (Pinyon-Juniper Ecosystem). K-21 (Juniper-Pinyon Woodland). Oak variant of SRM 504 (Juniper-Pinyon Pine Woodland). SAF 239 (Pinyon - Juniper). Oak-Pine Series123.32, oif Madrean Evergreen Forest and Woodland, 123.3 (Brown et al., 1998). Chihuhuan Deserts- Chihuhuan Montane Woodlands Ecoregion 24 d (Griffith et al., 2004).

283. Immature fruit of smooth sotol- A flowering-fruiting stalk of smooth sotol with an abundance of immature fruit. The Dasylirion species are dioecious so this was a pregnant girl plant. According to Powell (1988, p. 70) smooth sotol is the most common Dasylirion species in the Trans-Pecos Basin and Range area.

Presidio County, Texas. June.

284. Lechuguilla (Agave lechuguilla)- This is probably the most abundant Agave species in the Trans-Pecos Basin and Range area. In fact, lechuguilla "has one of the most extensive ranges of the agaves" (Gentry, 1982, p. 154). Lechuguilla is also one of the most easily identified (usually recognized immediately) by the high number of rosettes growing in clusters or groups. A. lechuguilla is a highly modular organism. Each of the basal rosettes is a module (a clone= ramet) of a genetic indivisual (genet). Most reproduction is asexual via suckering from rhizomes or rhizome-like structures (ie. "rootstocks"). Gentry (1982, p. 154) stated that numbers of rosettes "probably exceed those of all other native agaves". Gentry (1982, ps. 30-31) described the Agave rosette.

Incidentally this wonderful book, Agaves of Continental North America, (Gentry, 1982) is the encyclopedia and definitive reference by the man regarded as the world expert on Agave. Expensive, but well worth the price to any succulent-lover.

Lechuguilla is a poisonous range plant that has been documented to poison cattle, sheep, and goats, and--no surprise--this poisoning takes place under conditions of overgrazing. Gentry (1982, p. 157) delightfully described such mismanagement concluding that lechuguilla is "a protective agent of the range, penalizing those stockmen who, through force of circumstance or lack of foresight, decimate their resource by over-use". AMEN! The poisonous principle in lechuguilla is a saponin that causes hepatogenic or secondary photosensitization due to liver damage. References include Kingsbury (1964, ps. 56, 467-468), Sperry et al. (1964, ps. 7-8), Burrows and Tyrl 2001, 13-15), and Hart et al. (2003, 22-23).

Taxonomic treatment of Agave has been controversial. Traditionally the genus has been included in the agave family, Agavaceae, (eg. Powell, 1988), but other workers (eg. Smith, 1977) placed Agave in the lily family, Liliaceae, as an agave subfamily, Agavoideae. Incidentally, is there anything that is not controversial when it comes to plant taxonomic treatments?

Hudspeth County, Texas. June.

285. Basal rosettes of lechuguilla- Example of a cluster of rosettes, asexual modules of a genetic individual of lechuguilla. It was explained in the immediately preceding photo-caption that lechuguilla probably produces more rosettes than any other Agave species in North America. Recall from that explantion that each such rosette is a clone (= module= ramet) of the original "parent plant" which is the genet. Gentry (1982, p. 30) explained that each rosette of A. lechuguilla is a "monocarpic rosette". Each rosette flowers only once in its life (the life of that clone or module) and then dies; in fact, it begins to die as soon as the one-time flower stalk with its inflorescence begins to emerge and elongate. This condition was obvious in the flowering rosette in the cluster shown in this photograph. The inflorescence on this particular stalk was presented in the immediately succeding photograph.

With this pattern of life cycle and resource allocation, each genetic individual (the genet or actual unique plant) of lechuguilla with its ramets (rosettes) is a "multiannual" (Gentry, 1982, p. 30). Gentry's choice of terms could confuse the beginning student. Yes, the individual rosette flowers only once in its life and then promptly dies. The flowering-seed production process takes only a few weeks as flower stalk growth is extremely rapid (perhaps over a foot a day) such that this phenological development is "annual" (more like "ephemeral"). Yet it takes years (perhaps a quarter century) of reserve food storage, rosette growth, and formation of stalk primordial tissue development before flowering can be initiated. In reality each rosette is a long-lived perennial that finally flowers and then summarily dies. With asexual (vegetative or clonal) reproduction the genetically unique plant has a life span that is seemingly "endless" or "forever". The only thing annual about an Agave species is the amazing flowering and fruit production phenomenon. And that is phenomenonal.

288. Havard agave or Havard century plant (Agave havardiana)- Life and death simultaneously were playing out in this sexually reproducing Havard agave. After taking years (maybe several decades) to store up reserve food, grow leaves of the basal rosette, and development floral meristem this agave was in full-bloom and dying at the same time. The lowrmost leaves had already died as this plant was in anthesis and in process of exchanging gametes.

A. havardiana is in the subgenus Agave which has the paniculate form of inflorescence. A drooping or weeping juniper with fire-killed outer branches "watched" from behind as timeless Casa Grande provided a proper backdrop. Range Management has a romance all its own.

Chisos Basin, Big Bend National Park, Brewster County, Texas. June.

289. Mexican pinyon pine (Pinus cembroides)- Two specimens of Mexican pinyon pine growing in Green Gulch, Big Bend National Park (Brewster County, Texas). The "nut pines" have commonly been interpreted as one of four subdivisions of the soft pine group (subgenus Haploxylon; Strobus). Treatment by Harlow et al. (1979, ps. 60, 74-76) was readily followed. The so-called "nuts" are merely the naked seeds of a complicated taxon within Pinus. These seeds have served as important feeeds (ie. mast) for native and domestic animals as well food as for man. American Indians relied on pine nuts as a dietary stable for generations extending back to prehistory.

The Mexican pinyon pine of Trans-Pecos range environments is Pinus cembroides var. cembroides (Harlow et oal., 1979, p. 76; Powell, 1988, p. 51).

291. Trunk of alligator juniper (Juniperus deppeana)- The bark of alligator juniper is separated or divided into rectangular units that somewhat resemble the scales on the back of the American alligator (Alligator mississippiensis). This juniper (it is not usually referred to as "cedar" as are many of the species of this genus) is one of the largest and, with the tell-tale bark, most distinctive of the western Juniperus species. In most of the pinyon-juniper woodland subtypes of which it is a component alligator juniper is not the most juniper nor is it typically the only Juniperus species. With its large size, however, alligator juniper has usually been regarded as a co-dominant especially when it grows in association with one of the pinyon pines. The trunks of alligator specimens are often seen to be of odd shape or form in addition to relatively large size (as shown in this individual).

Green Gulch, Big Bend National Park, Brewst4r County, Texas. June.

292. Boughs (with cones) of alligator juniper- Branches with needles and fleshy cones of Juniperus deppeana. So-called "berries" on junipers are obviously not berries because the gymnosperms bear naked seeds. Seeds of Gymnospermae members are often borne together (in groups) on a woody structure known as a cone (eg. pine cone). In case of Juniperus species, the naked seeds are encased inside a fleshy exterior (hence "berry" to the laymen) that is termed a fleshy cone or, sometimes (and less properly), as fleshy seed.

Fleshy cones and scaley leaves (needles) were displayed in more detail in the second of these slides.Green Gulch, Big Bend National Park, Brewster County, Texas. June.

293. Time for a unique one- Arizona cypress (Cupressus arizonica) is the sole species of this genus that grows in the Trans-pecos Basin and Range Region. With Juniperus species, Cupressus arizonica rounds out representation of the cypress family, Cupressaceae. Arizona cypress ranges from the northern states of Mexico westward to California and as far north and east as New Mexico, but its only natural occurrence in Texas is in the Chisos Mountains of Brewster County.

Arizona cypress has much value as a native ornamental (and is grown far beyon its species range). Shown here was one small terminal branch (first slide) and mature though yet unopened cones (second slide).

Brewster County, Texas. Late October.

294. Cypress cones- Opening cones of Arizona cypress. The cones of Cupressus species are more like some members of the redwood or bald cypress family, Taxodiaceae, than they are like some members of Cupressacea such as Juniperus species which, as shown above, have fleshy cones. At maturity, cones of Arizona cypress dehisce or split open to release the dry seeds contained inside as seen in these two photographs..

Brewster County, Texas. Late October.

300. Emory oak (Quercus emoryi)- Leaves and acorns on a large Emory oak in the Davis Mountains of Trans-Pecos Texas. Emory oak is in the red or black oak subgenus, Erythrobalanus, species of which require two full growing seasons (years) for acorn production. Acorns are, of course, one of the major fruits known as mast, "nuts, acorns, and similar products which are consumed by animals" (Jacoby, 1989). Species from native birds and ruminants to man and his livestock use acorns as a staple of their diets.

Jeff Davis County, Texas. June.

301. Gray oak- Gray oak is the dominant species of scrub oak in most of the major mountain ranges throughout the Trans-Pecos Basin and Range area. This specimen was the plant growing to the right of the Emory oak tree in the two photographs just before the preceding photograph of Emory leaves and acorns.

Green Gulch, Big Bend National Park, Brewster County, Texas. June.

302. Leaves of gray oak (Quercus grisea)- Leadaers with leaves of the specimen of gray oak shown in the immediately preceding photograph.Gray oak is in the white oak subgenus, Leucobalanus, species of which require only one full growing season for production of an acorn crop.

Green Gulch, Big Bend National Park, Brewster County, Texas. June.

306. Baby leaves above baby girls- New leaves and young pistillate catkins of Lacy oak (Quercus lacyi= Q. glaucoides) against a blue spring sky.

Brewster County, Texas. Mid-March.

307. Leader of Lacy oak- With backdrop of bright autumn sky the two distal-most nodes (last two growing seasons of growth) of Lacy oak portrayed characteristic leaves plus some acorns.

Brewster County, Texas. Mid-September, mature-fruit stage.

308. Leaves and acorns of Lacy oak- Detail of leaves and acorns of Lacy oak. This species is in the Leucobalanus subgenus so nuts grow to maturity in a single year (one warm-growing season).

Brewster County, Texas. Mid-September, mature-fruit stage.

309. Lacy concentrate- Acorn of Lacy oak showing fruit characteristics of this species. The hgig-nutrient concentration of acorns qualifies this nut as a concentrate range feed.

Brewster County, Texas. Mid-September, mature-fruit stage.

310. Two "pals" in the oak-pinyon-juniper woodland cover type- A Texas madrone, known also as naked Indian and lady's leg, (Arbutus texana= A. xalapensis= A. xalapensis. var. texana) and, behind the madrone, alligator juniper. Texas madrone typically grows as an individual tree or shrub or, infrequently, in small colonies. This species is not a dominant or even an associate species, but it is a woody component and characteristic species of oak-pinyon-juniper woodland range in the Trans-Pecos Basin and Range area. Texas madrone (like alligator juniper) is also a characteristic species in range plant communities besides the pinyon-juniper cover type. An example includes ecotones between a mosaic of encinal oak and pinyon-juniper woodland and ponderosa pine cover types which was the case for these two trees. Texas madrone does "stand out" in the any woodland crowd as rangemen would expect of a lady's leg or a naked Indian.

Texas madrone is in Ericaceae, the heath or heather family, one of the characteristics of which is smooth, colorful bark on often bent or curving shoots and limbs (again, like a lady's leg or naked Indian).

Guadalupe Mountains, Guadalupe National Park, Culbertson County, Texas. June.

311. Two "legs" in the oak-pinyon-juniper woodland cover type- A "pin-up" photograph of Texas madrone or lady's leg (right) and an alligator juniper (left) showing characteristic bark of these two species that occur on oak-pinyon-juniper woodland range. Close-up of the two trees introduced in the immediately preceding photograph. These two specimens were growing in the Guadalupe Mountains in transition range vegetation between encinal oak/ pinyon-juniper woodland and ponderosa pine forest.

Leaves of ericaceous species are, like their bark, distinctive and possessing unique features. Leaves of most members of the Ericaceae are relatively large, thich, and highly cutinized. Ericaceous leaves such as those of Arbutus species have traditionally been described as "broad-schlerophyllus" following Cooper (1922) who first applied the term to Clifornia chaparral vegetation.

Guadalupe National Park, Culbertson County, Texas. June.

312. Branches of littleleaf sumac (Rhus microphylla) in full-fruit- Leaders of littleleaf or desert sumac (first photograph) and details of leaves and fruit of the same (second slide). The fleshy fruit of Rhus species is a drupe: the fruit type in which the outer layer or skin is the exocarp, the fleshy layers are mesocarp, and the bone-like seed coat is the endocarp; endocarp and seed constitute a pyrene (the stone or pit). Hence, drupes are stone fruits or pit fruits (Smith, 1977). This woody species is well-adapted to xeric environments and is widely distributed on harsher habitats such as drier or shallower soils in the Great Plains as well as basins and bajadas of the Chihuhuan Desert. Desert sumac is often common of tobosagrass (Hilaria mutica) swales or "flats" in the semidesert grasslands.

Drupes are sometimes cooked in hot water to make "hillbilly lemonade". Beats a job in the eye, but this author recommends the "real stuff" (or tea or coffee or milk or cold water or ...; you get the picture).

Davis Mountains, Brewster County, Texas. June.

314. Leaves and inflorescences of lanceleaf sumac- This Rhus species is of limited browse value for livestock (generally Poor feed value) except for goats and it is of somewhat higher browse value (roughly Fair) for native browsers like deer, but the drupe fruits can be excellent for birds like upland game species. All visible parts of this woody species appeared to this author to have potentially high ornamental value, especially for native plant purists.

The inflorescence type is a panicle.

Davis Mountains, Brewster County, Texas. June..

315. Evergreen sumac, tobacco sumac, or lentisco (Rhus virens subsp. virens)- Not much attention has been paid to this sumac as a range plant. In Important Western Browse Plants Dayton (1931, p. 97) cited previous work that found evergreen sumac "to be a pioneer in the vegetation of limestone ledges in west Texas, forming a protection for other species which occupy more slowly..." and therefore being important in watershed protection.

The adjective "evergreen" comes from the supposed characteristic of evergreenness, but leaves of this subspecies and toughleaf sumac (R. virens subsp. choriophylla) frequently turn red or brown in autumn. Leaves are extrmely shiny and bright. The adjective "tobacco" was derived from reports that Indians like the Commanches cured leaves and used them as a substitute for or mixed with tobacco to smoke in their pipes. The fruits of all Rhus species can be used to make a refreshing drink (often termed "lemonade") by steeping in hot water and subsequent cooling. Good for college freshmen who will (do) try any- and everything. Returning to the namesake "tobacco" and sticking with college students, it should be pointed out that smoking of Rhus leaves is likely less toxic than most else they smoke, snort, guzzle, shoot, poke, prod, pump, hump, or otherwise imbibe in.

Presidio County, Texas. June.

316. Phytoparasite in the Chisos- Rough or boll American mistletoe (Phoradendron bolleanum= P. hawksworthii) parasitizing a juniper (Juniperus sp.). Add to the multi-layers of the oak-Mexican pinyon pine-juniper woodland an epiphyte layer, and a parasitic one at that. Parasitism is one of the important interactions among organisms on ranges. In context of range plants and vegetation your author included important parasitic throughout this publication. This example continued that tradition.

Mistletoes have been extensively studied by forest pathologists, but the mistletoe featured here is not one of the major economically important species.

Mistletoes like the Phoradendron species are known as the true or leafy mistletoes (in contrast to the dwarf mistletoes that are far more common on conifers).

Green Gulch, Big Bend National Park, Brewster County, Texas. June.

317. Plant parasitism in the Chisos- Rough or boll American mistletoe parasitized (and probably killed) this juniper on an oak-Mexican pinyon pine-juniper woodland range. The second slide showed the sites of shoot emergence and of penetration of haustorium which is the projection of hyphae that function as the absorbing organ into tree cells. Haustorial strands spread laterally along conifer shoots (branches) and send down sinkers deeper into shoots to absorb nutrients of the host conifer (Agrios, 1988, ps. 618-620).

Green Gulch, Big Bend National Park, Brewster County, Texas. June.

318. Chinograss or chino grama (Boutelous ramosa)- This is the dominant range plant on much of the foothill semidesert grassland and Chihuhuan Desert-semidesert (Chihuhuan) grassland ecotone range types and it is an important (often co-dominant) species in the sotol-lechuguilla-grama grassland range type. Chino grama remains a major species in the herbaceous layer in the oak-Mexican pinyon pine-juniper woodland cover type where it is typically the most widespread grass. In the oak-pinyon pine-juniper woodland individual plants of chino grama attain considerably less size, especially fewer shoots, than on grassland or shrub-savanna range types.

The rocky land surface shown here was typical of much of the oak-Mexican pinyon pine-juniper woodland. There were not relict or reference areas to which this soil surface could be compared. This surface condition was very similar to stone pavement on floor of the bajadas and basins of Chihuhuan Desert below the foothills of Chisos Mountains where this photograph was taken.

Green Gulch, Big Bend National Park, Brewster County, Texas. June, early estival stage prior to onset of summer rainy period and, therefore, prior to this year's "green-up".

319 Colony of spidergrass or spider threeawn (Aristida ternipes= A. trnipes var. ternipes)- This is one of the more distinctive (ie. less easily confused) of the perennial Aristida species. It is also one of the more abundant threeawns in mountains of the Trans-Pecos area and, indeed, throughout much of the immense Chihuhuan-Sonoran Deserts super-region. While spidergrass is distinctive (panicles were shown below), and not as readily confused as most of the semiarid-arid Aristida species, this does not preclude mix-ups, nomenclatural inconsistency, continuing analysis and re-analysis of specimens, and spirited contradictions in taxonomy of A. ternipes (along with most other Aristida speceis).

Some of the confusion in the literature on Aristida was described in detail elsewhere herein, especially in regards the numerous Aristida species in the Sonoran Desert chapter (Shurblands), so details were not "re-hashed" in this photo-caption. Various authorities who tried to "straighten out the mess" (and wound up contributing to the mess as well as to Aristida taxonomy) in southwestern North America included Silveus (1933), Hitchcock and Chase (1951), Gould (1951), Gould (1975), and Powell (2000). As of this writing the latest "official" treatment (but if the past is any indication not the last of such) on Aristida was that of Allred (in Barkworth et al., 2003, ps. 330-335). Also worthy of reference was the well-structured work of Powell (2000, ps. 239-254 passim, esp. 244-246).

The latest taxonomic treatments viewed A. ternipes and A. hamulosa as a "species pair" and "cospecific" as A. ternipes var. ternipes and A. ternipes var. hamulosa (see Powell, 2000, ps. 245-246 for discussion and citation of original sources).

Specimens of spidergrass presented in this and the next two slides were representatives of the dominant herbaceous species in the understorey of upper elevation oak-Mexican pinyon pine-juniper woodland range in the Chisos Mountains.

Green Gulch, Big Bend National Park, Brewster County, Texas. June, seed-ripe to seed shatter phenological stages.

320. Specimen of spidergrass- This individual of Aristida ternipes was a good example of the habit (general appearance) and some of the gross physical details of its species. The tortuous shape of upper leaves and panicle branches could have easily invoked metaphorical comparison to a spider-like configuration.

Green Gulch, Big Bend National Park, Brewster County, Texas. June, seed-ripe to seed-shatter stage.