Forests and Woodlands - Range Types of North America

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 pinandhills and through the mixed prairie in northwestern Okahoma. e 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 S

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.

1. The Black Hills ecosystem (or landscape mosaic) includes range types that go from the tallgrass and mixed prairies (covered above) to the ponderosa pine and ponderosa pine-bur oak-paper birch (Betula papyrifera) forest types. This overall view of the Black Hills is primarily ponderosa pine forest with the Pre-Cambrian uplifted granite domes in the central area. Custer State Park, South Dakota. July. FRES No. 21 (Ponderosa Pine Ecosystem). K-16 (Black Hills Ponderosa Pine Forest).SAF 237 (Interior Ponderosa Pine). No SRM for the plains ponderosa pine type. Middle Rockies- Black Hills Plateau Ecoregion, 17b (Bryce et al., undated).

2. Panoramic scene of the Black Hills and pristine ponderosa pine-Black Hills National Forest, South Dakota. July. FRES No. 21 (Ponderosa Pine Ecosystem). K-16 (Black Hills Ponderosa Pine Forest). SAF 237 (Interior Ponderosa Pine); no SRM. Middle Rockies- Black Hills Plateau Ecoregion, 17b (Bryce et al., undated).

3. Interior of the ponderosa pine cover type with paper birch, Black Hills or white spruce (Picea glauca) and bur oak forming an open forest with a classic prairie understory dominated by little bluestem and prairie dropseed with the introduced but naturalized European grasses, Kentucky bluegrass (Poa pratensis ), timothy (Phleum pratensis), smooth bromegrass, and orchardgrass (Dactylis glomerata). Black Hills National Forest, Pennington County, South Dakota. July. FRES No. 21 (Ponderosa Pine Ecosystem). K-16 (Black Hills Ponderosa Pine Forest). SAF 237 (Interior Ponderosa Pine); no SRM. Middle Rockies- Black Hills Plateau Ecoregion, 17b (Bryce et al., undated).

4. Understory of ponderosa pine-bur oak-eastern cottonwood forest type supporting western wheatgrass, slender wheatgrass, little bluestem, prairie dropseed, bluegrasses and needlegrasses (including Stipa richardsonii and S. occidentalis ). Silty foot slope range site. Custer State Park, South Dakota. July. FRES No. 21 (Ponderosa Pine Ecosystem). K-17 (Black Hills Ponderosa Pine Forest). SAF 237 (Interior Ponderosa Pine); no SRM. Middle Rockies- Black Hills Plateau Ecoregion, 17b (Bryce et al., undated).

5. Composite shot of ponderosa pine range with paper birch and quaking aspen as associates of the dominant pine. Understory is comprised primarily of the introduced festucoid grasses, Kentucky bluegrass, smooth brome, and timothy. Custer State Park, South Dakota. July. FRES No. 21 (Ponderosa Pine Ecosystem). K-17 (Black Hills Ponderosa Pine Forest). SAF 237 (Interior Ponderosa Pine); no SRM. Middle Rockies- Black Hills Plateau Ecoregion, 17b (Bryce et al., undated).

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).

Rocky mountain-Great Plains Transition

The following section presented a ponderosa pine forest and forest-grassland (true prairie) transition on the western margins of the Northern Great Plains. This section was also included with the chapter, True Prairie, because it was a natural ecotone between true prairie and and an outlier of the eastern ponderosa pine type (SAF 237). This natural range plant community was included in this chapter as well (and not in the chapter devoted to forests range types of Northern Rocky Mountains) because this range vegetation was an "island" of the ponderosa pine forest cover type( SAF 237) that develops in the continental interior of North America.

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).

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.

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.

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.

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.

14. Spikes 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.

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.

15. Two biomes, ecosystems, and range types in one- Ponderosa pine forest with glade (natural opening of herbaceous vegetation) of western wheatgrass (Agropyron smithii), green needlegrass (Stipa viridula), and needle-and-thread (S. comata) in the Northern Great Plains or, more specifically the eastern unglaciated or sedimentary plains portion of the Missouri Plateau.

Inside the forest (ie. under the ponderosa pine canopy) the understorey was similar to that of the glades except that in more mesic areas beneath the pines and near a small streeam there was more cover of Kentucky bluegrass (Poa pratensis), a naturalized, Eurasian, rhizomatous, perennial grass. Kentucky bluegrass was absent in the natural grassland of the glades. For all practical purposes there were no forbs other than a few robust plants of the naturalized, Eurasian weed, hound'stongue (Cynoglossum officinale).

These native, perennial grasses were ranked as increasers on this range site (Ross and Hunter, 1976, p. 16) such that this forest range was in high Fair to low Good range consition class. Bluebunch wheatgrass, Idaho fescue, little bluestem (Andropogon scoparius), and sideoats grama (Bouteloua curtipendula) were listed as climax dominants (ie. decreasers) for this range site (Ross and Hunter, 1976, p. 16).

"For all intents and purposes" there were two possible interpretations of this range vegetation. One interpretation was that this natural pasture consisted of two forms: 1) a forest range of wheatgrass-needlegrass true prairie or mixed prairie with ponderosa pine and 2) a grassland range of wheatgrass-needlegrass prairie without trees. A second interpretation was that this was a single range plant community, being a ponderosa pine--western wheatgrass--needle-and-thread--green needlegrass forest with some glades (natural openings) that occurred amid a more-or-less closed canopy forest.

"Lumpers" and "splitters" exist among vegetation scientists as well as taxonomists.

Musselshell County, Montana. Mid-June; vernal aspect. FRES No, 21 (Ponderosa Pine Forest & Woodland Ecosystem. K-15 (Eastern Ponderosa Pine Forest. SAF 237 (Interior Ponderosa Pine). Cold Temperate Forest and Woodland 122-Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62, Pinus ponderosa Assocaition 122.621 of Brown et al, (1998, p. 37). FRES No. 38 (Plains Grassland Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 607 (Wheatgrass-Needlegrass). Nothing appropriate in Brown et al. (1998). Eastern Sedimentary Plains- Forest-Grassland Complex range site, 15-19 inch precipitation zone, (Ross and Hunter, 1976, ps. 16-17). Northern Great Plains, Pine Scoria Hills, Ecoregion 43p (Woods et al., 2002).

16. Outside and inside; two ecosystems (or, is it, three?)- Exterior of a ponderosa pine--western wheatgrass--green needlegrass--needle-and-thread range plant community in background and a (first slide) and interior of the ponderosa pine western wheatgrass-needlegrass forest range (second slide). This range vegetation was in the unglaciated or eastern sedimentry plains portion of the Northern Great Plains in central Montana.

The range plant community in foreground of the first slide was riparian vegetation growing along a small stream. This streamside vegetation consisted mostly of western snowberry (Symphoricarpos occidentalis) with some naturalized Kentucky bluegrass. This hydric vegetation could be regarded as comprising a different ecosystem from the forest range ecosystem in the background of this first slide and the range plant community of the second slide.

Also visible in the second slide was a natural opening or glade in the ponderosa pine forest. Although the herbage of the tree-free glade and that in the understorey below the pines was the same (western wheatgrass, needle-and-thread, and green needlegrass), the glade community and the pine-wheatgrass-needlegrass community could be seen as two different range communities or, alternatively, as one forest range community with some grassy openings.

17.Trees, grass, and cow brutes- Interior of a ponderosa pine forest with an understorey dominated by western wheatgrass, green needlegrass, and needle-and-thread that was a range for cow-calf pairs. This forest range was in the eastern sedimentary or unglaciated part of the Northern Great Plains in central Montana.

18. Ground-level of an ecotone- Understorey of a ponderosa pine--western wheatgrass--green needlegrass--needle-and-thread forest range at outer edge of a riparian zone dominated by western snowberry. The broadleafed plants were clonal offshoots of the strongly rhizomatous western snowberry. Most of the herbadeous turf was Kentucky bluegrass. The cow chip in lower lkeft foreground attested to presence of brood cows on this range and reminded students of nutrient recycling in range ecosystems.

19. Different breed of dog- Two "ramboucous" plants of hound'stongue (Cynoglossum officinale) presented in the first or upper slide with the lower or second slide showed the upper part of plant with its inflorescence. This member of the borage family (Boraginaceae) is a native of Eurasia that, after being brought over by white man, decided that it like it almost as well as he did. This rank-growing, forb is a biennial that has naturalized over much of North America. Houndstongue is best regarded as weed with its preferred habitats being "disturbed pastures, roadsides, forest edges and meadows" (Great Plains Flora Association, 1986). This one was growing just beyond a borrrow ditch along an old logging road beside a ponderosa pine-abunchgrass range.

For treatment as a weed, Weeds of the West (Whitson et al., 1992, ps. 202-203) was recommended. This treatment described briefly the poisonous nature of hound'stongue with the poisonous principle being pyrrolizidine alkaloids tht cause cessation of liver regeneration. Of course, the encyclopedic Toxic Plants of North America (Burrows and Tyrl (2013, ps.269-272) provided ultimate treatment of this poisonous plant and its toxxicity. Burrows and Tyrl (2013, p. 270) specified that the pyrrolizidine alkaloids involved are ester forms of heliotridine and supinidine. Stock poisoning is commonly fatal, plus the pyrrolizidine alkaloids pose a threat to hman health because they are potential corcinogens that are sometimes found at low concentrations in milk and liver (Burrows and Tyrl, 2013, p. 272)

20. Head and tongue- Sexual shoot with flower clusters (upper slide) and flower (lower slide) of hound'stongue growing on a ponderosa pinebunchgrass range in the Unglaciated Missouri Plateau. The inflorescence of this eye-catching forb is a thyrse collectively formed from numerous cymes (Great Plains Flora Association, 1986). Cyme refers to "a general inflorescence type characterized by flowers whcch bloom from the center outward or the apex downward" while thyrse is "a condensed panicle-like inflorescence in which the main axis is indeterminate and the lateral branches are determinate" (Smith 1977, ps. 293, 310). In other word,s this complicated inflorescence is an inflorescence (flower cluster) within an inflorescence or an inflorescence composed of inflorescences.

All in all this weed is a unique, introduced-and--staying-for-the-remainder member of various flora, including in this case one of the many variants of the Pinus ponderosa range cover type.

In the following section two examples of the savanna form of the ponderosa pine-bunchgrass range cover type that had developed in the sedimentary hills above the Townsend Basin in southern Montana.

Ponderosa pine forms range vegetation that varies from relatively dense forest to open forest through to woodland and, finally, savanna. The herbaceous understorery of these various forms may be the same with the only substantial difference being in tree cover.

21. Some trees and a lot of grass- A savanna of ponderosa pine with an herbaceous understorey (no lower woody layer as no shrubs were preent) dominated by bluebunch wheatgrass and with green needlaegrass and slender wheatgrass (Agropyron trachycaulum= A. subsecundum in the treatment by Lesica [2012, p. 657]) as associate species. The major forbs were two naturalized, Eurasian, biennial, weeds, flannel mullein (Verbascum thapsus) and common or bull thistle (Cirsium vulgare). There were other forbs at other local microsites in this ponderosa pine-bunchgrass savanna. These were native forbs and they were shown below.

Logging trucks had traveled over this immediate area (near a road right -of-way) three summers prior to photographs and disturbance to the soil surface permitted growth of these pioneering, weedy forbs. This local disturbance and the dead stalks of flannel mullein were visible in the first of these two slides. Logging of ponderosa pine had not been been on this immediate area, but logging truck activity had occurred at edge of this savanna.

Pines in this savanna were all second-growth trees, complete with a few pine seedlings and saplings.

This natural vegetation (except for a few naturalized, pioneering weedy, forbs) could be regarded as either a ponderosa pine forest of widely scattered trees with a wheatgrass-needlegrass understorey or, alternatively, as a wheatgrsss-needlegrass grassland with a sparse cover of ponderosa pine.

Lewis & Clark County, Montana. Late June; estival aspect. There were two options regarding classification of this range vegetation. FRES FRES Middle Rockies- Townsend-Horseshoe-London Sedimentary HillsFRES No, 21 (Ponderosa Pine Forest & Woodland Ecosystem. K-15 (Eastern Ponderosa Pine Forest. SAF 237 (Interior Ponderosa Pine). Cold Temperate Forest and Woodland 122-Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62, Pinus ponderosa Assocaition 122.621 of Brown et al, (1998, p. 37). FRES No. 38 (Plains Grassland Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 607 (Wheatgrass-Needlegrass). Nothing appropriate in Brown et al. (1998).Foothills & Mountains- Forest-Grassland Complex, 15-19 inch precipitation zone (Ross and Hunter, 1976, p. 30). Townsend-Horseshoe-London Sedimentary Hills Ecoregion 17y (Woods et al., 2002).

22. Trees were hit-and-miss, but grass was always there- A savanna of apparently young, second-growth ponderosa pine with an herbaceous layer dominated by bluebunch wheatgrass with slender wheatgrass and green needlegrass as associate species.

23. Little trees and big grass- Comparatively large up to huge sized plants of bluebunch wheatgrass (dominant herbaceous species), slender wheatgrass, and green needlegrass (these two were associate herbaceous species) on a ponderosa pine savanna on sedimentary hills above the Townsend Basin in southern Montana. All three grass species were abundant in the first of these two slides whereas bluebunch wheatgrass was the only species visible in the second slide. Specimens of bluebunch wheatgrass seen in the second slide were extremely large plants with a pronounced cespitose (bunched= clumped; bunchgrass) habit.

A few trees had been harvested near a road right-of-way on close proximity to the foreground of the area presented in the second slide. Tree removal could have reduced tree-grass competition as a major factor in development of bluebunch wheatgrass to gigantic dimensions (over two and a half feet in height and two feet across foliar cover). Some of these giant bluebunch wheatgrass plants were presented in the immediately succeeding slide.

24. A big herbaceous specimen- Three relatively huge plants of bluebunch wheatgrass (two flanking and slightly to the rear of the centermost plant) on a savanna of ponderosa pine, bluebunch wheatgrass, green needlegrass, and slender wheatgrass that development on sedimentary hills above Townsend Basin in southern Montana.

In the local microhabitat seen here (immediate edge of a road right-of-way) a young ponderosa pine (roughly 16 inches diameter breast height) had been felled and taken to the mill. Harvest of this tree might have contributed to the rank-growth and immense size attained by these specimens of bluebunch wheatgrass; however, there were other plants of bluebunch wheatgrass (and of associate species, slender wheatgrass and green needlegrass) growing under ponderosa pine in a savanna form that were almost as large. (These specimens were simply too impressive size-wise to not include in this discussion even if an Epson Perfectiont 700 scanner over-scanned them to some degree.) These examples attested to the large size and rank-growth habit of which bluebunch wheatgrass is capable.

No livestock were on this range so that any grazing by large herbivores would been that of deer. The straw or dead shoots in these bunchgrasses were last year's grassstalks. Prescribed fire should have been or, at least, could have been a desirable practice on this forest (savanna) range.

25. Forb part of the herbaceous layer- Local microsite in the herbaceous layer of a ponderosa pine-bluebunch wheatgrass-green needlegrass-slender wheatgrass savanna in the sedimentary hills above the Townsend Basin in southern Montana. Plant species in this "photo-quadrant" included bitterroot (Lewisia rediviva); State Flower of Montana (and in early bloom stage); lesser, Rocky Mountain, or prairie spike-moss, (Selaginella densa= S. rupestris var. densa= S. scopulorum= S. standleyi); and, at pre-bloom stage, dwarf or yellow mentzelia; desert or golden blazing star; bullet, or dwarf, or yellow stickleaf (Mentzelia pumila).

There were several such local microhabitts on this savanna range, the exact cause of which was unknown to this author. They appeared to be local disturbances, but the source of such disturbances was not obvious.

26. One slope had pines and grass; another slope had only grass- An example of a ponderosa pine-bluebunch wheatgrass--needle-and-thread--Sandberg's bluegrass savanna in the sedimentary hills above Townsend Basin in central Montana. Bluebunch wheatgrass was the herbaceous dominant (actually, the dominant range plant of this vegetation overall) while needle-and-thread and Sandberg's bluegrass were its associates. Gravel, woolypod, or Pursh's milkvetch (Astragalus purshii var. concinnus) was the only forb of consequence on this savanna range that was obviously in Excellent range condition class. An indicator of the ner-virgin state of this range was the near absence of cheatgrass or downy brome. Clearly the climax range vegetation of this range site.

While the overall range vegetation in this example was that of a savanna (savanna form of ponderosa pine-bunchgrass understorey) there were local stands of ponderosa pine of such density and dispersion as to comprise a woodland (trees were closer than in a savanna but their crowns did not touch each other or interlock) while other areas were bludbunch wheatgrass-dominated brassland (ie. a bunchgrass steppe).

More typical savanna physiogonomy was presented in the next two slide-caption units.

27. Grass was more at home than the trees- A ponderosa pine-bunchgrass savanna in sedimdntary hills above Townsend Basin in central Montana. The dominant plant species in this range plant community was bllebunch wheatgrass. Assolciagte grass species were needla-and-thread and Sandberg's bluegrass. The only forb of mentionable cover was gravel, woolypod, or Pursh's milkvetch. Downy brome or cheatgrass was--for all intents and purposes--combpletely absent which bespoke of the near-virgin state of this relict of climax range vegetation.

Yes, an Epson Perfection 700 scanner fouled up the second slide, and even Adobe PhotoShop could not restore it to the original color of Provia 100f film. Never buy anything Epson.

28. Savanna in the hills- Range vegetation of a ponderosa pine-bunchgrass savanna that was an example of the ponderosa pine-bluebunch wheatgrass habitat type using the method of Daubenmire (1952, 1984). On this steep slope skunkbush sumac (Rhus trilobata= R. aromatica) contributed a shrub component though this was not of such cover as to comprise a shrub layer. Other important grass species included needle-and-thread and Sandberg's bluegrass. There were a few plants of gravel, woolypod, or Pursh's milkvetch in this climax range plant community. As an indicator of the pristine state of this range (Excellent range condition class) there was almost no cheatgrass or downy brome present.

29. Local sward of an associate species and probably a co-occurring species- Local population of Sandberg's bluegrass that was an associate species along with plains bluegrass (Poa arida), this latter perhaps a colony, on a ponderoda pine-bunchgrass savanna (a pondereosa pine-bluebunch wheatgrass habitat type).

Agrostologists have had a time (maybe a fun time, but cerainly a productive time from standpoint of peer -reviewed publications) debating the appropriate scientific name and related species, subspecies, varieties, whatever of the Poa secunda complex. Sandberg's bluegrass is one of several closely related Poa taxa. These were reinterpreted in the encyclopedic Flora of North America (Barkworth et al., 2007. ps. 586-588). This same interpretation was followed in Manual of Montana Vascular Plants (Lesica, 2012, p. 703). The typical Sandberg's bluegrass in this "neck of the woods" was described as P. secunda subsp. secunda.

Traditionally this taxon, with the common name of Sandberg's bluegrass, was known as P. sandbergii up to five or six decades ago, and since then as P. secunda. Sandberg's bluegrass was regarded by all authorities (eg. Skinner et al., 1999, p. 97) as being strictly tufted or cespitose (ie. having only tillers or intravaginated shoots) or, in words of Hitchcock and Chase (1950, p. 134), as having "a dense, often extensive, tuft of short basal foliage". For purposes of the International Intercollegiate Range Plant Identification Contest (Stubbendieck et al., 1992, p. 179) described P. secunda as "strongly cespitose". The Range Plant Handbook (Forest Service, 1940, G106) specified that Sandberg's bluegrass was "strictly a bunchgrass" that "…does not have underground stems (rhizomes, or rootstocks)…". (But read contracting details in the next parargraph.)

Sometimes tufted plants of Sandberg bluegrass grow so close together that they appear to form a more or less continuous sod. Such was the situation shown in this slide in which several plants of Sandberg's bluegrass grew in close spatial association and formed a sod-like sward. Most of the actual sod in this "photoquadrant", however, was cover of the rhizomatous plains bluegrass. Lesica (2012, p. 700) explained that "Poa arida is similar to and co-occurs with Poa secunda. Both have a bunched habit (albeit loosely in Poa arida), long narrow inflorescences, and a predilection to open, dry seetings". Such was the phenomenon seen by students in this slide. Furthermore, in some montane habitats P. secuda does have rhizomes and there is perhaps hybridization among some related Poa species (Lesica, 2012, p. 703).

30. Slender, awned, and/or bearded (take your choice) wheatgrass - Two plants of bearded, awned, or slender wheatgrass (Agropyron trachycaulum= A. subsecundum= just about everything else as described in the next caption) in understorey of a ponderoda pine forest (actually a savanna form)-bunchgrass range in foothills of Middle Rockies. These were robust plants exceeding two and a half feet in height.

Successional status of this slender/bearded wheatgrass taxon is apparently very range site-specific. Stubbendieck et al. (1992, p. 205) gave the forage value of slender wheatgrass as Excellent for sheep and cattle when green and still Good when herbage was mature. The Forest Service (1940, G7) described this grass as "a valuable forage lant, highly palatable to all classes of livestock". With a high palatability and forage value pretty much "nailed down" it would appear that slender or bearded wheatgrass would quickly decline with improper grazing management, but the Forest Service (1940, G7) continued that this species " is fairly resistant to grazing". In short, a damn fine range grass.

Then Lesica (2912, p. 657) had to go and add still more confusion to this embattled species by noting that slender (or bearded, awned, etc.) wheatgrass is "ecologically confined to disturbance-prone settings".

Even more confusing than response to disturbance is the name(s) of this valuable range species. This grass has about as confusing a name--common or scientific--as any plant species alive (see immediately succeeding caption).

Lewis & Clark County, Montana. Late June (summer solstice); early bloom stage of phenology).

31. Obviously bearded- Portions of two mature spikes of slender, awned or bearded wheatgrass in understorey of a ponderosa pine-bunchgrass savanna range Spikelets in these two spikes were fully mature with ripe grains and ready to shed them. As is the cse for most Agropyron species spikelets close down following pronounced exertion of anthers at anthesis. These two inflorescences were on plants growing in close proximity to the two cespitose plants presented in the immediately preceding slide.

This taxon (or taxa) is (are) one (some) of the most confusing and controversial of the cool-season grasses, especially those in tribe Tritaceae or Hordeae. Hitchcock and Chase (1950, ps.) distinguished between Agropyron trachycaulum and A. subsecundum. More recently in Manual of Montana Vascular Plants, Lesica (2012, ps. 656-657) used A. trachycaulum as the proper species and listed A. subsecundum as a synonym. Lesica (2012, p. 657) explained that A. trachycaulum var. unilaterale was a tall form of slender wheatgrass with lemmas having most awns as long as 40mm. For use in the Society for Range Management-sponsored International Intercollegiate Range Plant Identification Contest, Stubbbendieck et al. (1992, ps. 205, 459) also stayed with A. trachycaulum and listed A. subsecundum as a synonym. Stubbbendieck et al. (1992, ps. 205) specified that specimens of A. trachycaulum could have lemmas that were awnless or awned with awns up to 30mm long.

In a cladistic-based taxonomy Barkworth et al. (2007, ps. 321-324) moved slender wheatgrass to the genus, Elymus and distinguished between E. trachycaulus subsp. subsecundus, designated as one-sided wheatgrass, and E. trachycaulus sub. trachycaulus, designated as slender wheatgrass. Lesica (2012, p. 657) gave Elymus trachycaulus as a synonym, but retained the traditional, historical genus of Agroypron (thank goodness some taxonomists still have some sense about them). In their Flora of the Pacific Northwest (Hitchcock and Cronquist, 1973, ps. 614-615) used A. caninum subsp. caninum for bearded wheatgrass, with lots of varieties including var. majus (which had numerous synonyms including A. trachycaulum) for the common name of slender wheatgrass which had lemma awns up to 30mm in length. The Great Plains Flora Association (1986, p. 1124) used A. caninum subsp. majus (including variety unilaterale) for slender wheatgrass instead of A. trachycaulum, but Barkworth et al. (2007, p. 323) specified that A. caninum was restricted to Greenland and Eurasia.

Once again it seemed that common names have more continuity than scientific names (contrary to claims by herbarium taxonomists), but even here Hitchcockk and Chase (1950, p. 238) fed into the controversy and confusion early on when they used bearded wheatgrass for A. subsecundum and slender wheatgrass for A. trachycaulum!

Lewis & Clark County, Montana. Late June (summer solstice); grain-ripe phenological stage.

32. A wooly one- Woolypod, gravel, or Pursh's milkvetch (Astragalus purshii var. concinnus) in an open area within a ponderosa pine-bunchgrass savanna in the Middle Rocky Mountains of westcentral Montana.The genus Astragulus, commonly known as milkvetches or locoweeds (some of the latter are in other genera) comprises a species-rich (and very colorful) group of range plants. For example, in Montana Astragalus is the largest genus in the Leguminosae (legume family) with 49 species (Lesica, 2012, ps. 291-305). These are papilionaeous or the nodulated legumes and must be assumed to be valuable nitrogen-fixers.

From the perspective of Range Management, the most important aspect of Astragalus is their livestock-poisoning property. The poisonous plant feature of Astragalus species is a detailed field study unto itself. Students were referred to the classic poisonous plant literautre, most recently the comprehensive, encyclopedic treatise of Burrows and Tyrl (2013; ps. 502-526). The three main forms of livestock poisoning problems due to consumption of Astragalus spp. are locoism ("locaoweed poisoning") nitrotoxicosis, and selenosis (slelenium toxicity). Numbers of Astragalus species shown or strongly implicated as causing or contributing ot these three conditions were 38, 27, and 19, respectively Burrows and Tyrl (2013; ps. 507, 516, 521, tables therein). Some of these Astragalus species can cause two of the three forms of toxicity.

The showy woolypod or Pursh's milkvetch (and most Astragalus species are colorful forbs) causes only one of these toxicity problems: locoism. Woolypod milkvetch is apparently not one of the major locoweeds from standpoint of livestock losses (Burrows and Tyrl, 2013; ps. 502-526), but it is one of the more widespread--and therfore one of the better known --Astragalus species (Hermann, 1966, 58-60).

Lewis & Clark County, Montana. Late June; both peak-bloom and young legumes stages of phenology.

33. Wooly tops- Upper shoot with flowers and legumes (first slide) and details of papilionaceous flowers (second or lower slide) of woolypod or Pursh's milkvetch, one of the locoweeds that has been documented to induce locoism in livestock. The poisonous principle (toxic chemical) in locoism is a group of indolizidine alkaloids, such as swainsonine, produced br fungal endophytes (Burrows and Tyrl, 2013, ps. 509-512).

This plant was growing on a ponderosa pine-bunchgrass forest (technically, a savanna) range in the Middle Rocky Mountains of westcentral Montana.

Lewis & Clark County, Montana. Late June; both peak-bloom and young legumes stages of phenology.

34. Wooly blooms and legumes- Inflorescence and fruit (first slide) and telltale woody legumes of woolypod milkvetch produced on a bunchgrass-ponderosa pine savanna in Middle Rocky Mountains in westcentral Montana.

Lewis & Clark County, Montana. Late June; both peak-bloom and young legumes stages of phenology.

35. Understorey composite- Lava aster (Aster scopulorum) contently growing in the understorey of a ponderosa pine-bluebunch wheatgrass savanna in the foothills of the Middle Rocky Mountains.

36. Purty one in the pines- Nice specimen of dwarf or yellow stickleaf; dwarf or yellow mentzelia; desert or golden blazing star; bullet mentzelia, or (Mentzelia pumila) in the bluebunch wheatgrass-dominated herbaceous understorey of a ponderosa pine-bunchgrass savanna in the sedimentary hills above Townsend Basin in central Montana.

The common name of stickleaf comes from the tendency of leaves of the Mentzelia species to attach (stick) themselves to surfaces like fur and clothing. The Mentzelia species can be a major source of contamination in wool.

37. Showy parts- Deeply serrated leaves and the prominent inflorescence of dwarf, bullet, or yellow stickleaf in the bluebunch wheatgrass-dominated understorey of a ponderosa pine-bunchgrass savanna.

In spite of their showy appearance, Mentzelia species are not major range forbs. For instance, they did not merit coverage in the classics, Range Plant Handbook (Forest Service, 1940) or Notes on Western Forbs (Dayton, 1960; Hermann, 1966).

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.

38. "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).

39. 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).

40. 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.

41. 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).

42. 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.

43. 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).

44. 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.

45. 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).

46. 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.

47. 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.

48. 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).

49. 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.

50. Southwestern form of the ponderosa pine range type with uneven age population of pine— Open herbaceous understory of various sedges (Carex spp.) and in larger openings diverse grass component, including slender wheatgrass, mountain muhly (Muhlenbergia montana), pine or hairy dropseed (Blepharoneuron tricholepis), Arizona fescue (Festuca arizonica), muttongrass (Poa fendleriana), and bottlebrush squirreltail. Few forbs or shrubs but some Gambel oak.

Lincoln National Forest, Lincoln County, New Mexico. June. FRES No. 21 (Ponderosa Pine Ecosystem). Mapped by Kuchler as K- 17 (Ponderosa Pine-Douglas-fir Forest). SAF 237 (Interior Ponderosa Pine). No SRM for ponderosa pine cover types in the southwest region. Pinus ponderosa Association in Yellow Pine Series of Brown et al. (1998). Sacramento Mountains; Arizona/New Mexico Mountains- Rocky Mountain Conifer Forests Ecoregion, 23f (Omernik and Griffith, 2006).

51. 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.

Seen here is the upper reaches or montane zone of beautiful McKittrick Canyon, commonly regarded by connoisseurs of things Texas as the most scenic spot in the Lone Star State. It is the ponderosa pine montane forest in an arid zone. The striking Texas madrone known also as lady's leg and naked Indian (Arbutus xalapensis)) is the major associate on drier sites. One is seen here at left foreground. Bigtooth maple (Acer grandidentatum) assumes this role on more mesic sites. At still higher elevations quaking aspen occurs, including here in McKittrick Canyon, but it is not visible in this scene. Dominant shrub monocots include sotol (Dasylirion leiophyllum), the two plants in the foreground, and three species of agave: 1) New Mexico agave (Agave neomexicana]), 2) Havard agave (A. havardiana), and 3) slimfooted agave (A. gracilipes). Dominant grasses are sleepygrass (Stipa robusta), bull muhly or bullgrass (Muhlernbergia emersleyi), pine muhly (M. dubia), blue threeawn (Aristida glauca) and Wooton threeawn (A. pansa).

Guadeloupe Mountains National Park, Culberson County, Texas. Estival aspect, June. FRES No. 21 (Ponderosa Pine), K-Western Ponderosa Pine), SAF 237 (Interior Ponderosa Pine); no SRM for ponderosa pine cover type(s) in the Southwest. Pinus ponderosa Association in Yellow Pine Series of Brown et al. (1998). Arizona/New Mexico Mountains- Montane Woodlands Ecoregion, 23b (Griffith et al., 2004).

Location note and an introduction: treated in this section was a savannah or, at greatest tree density that one's imagination allowed, a woodland form of ponderosa pine-mixed prairie in the Davis Mountains of Trans-Pecos Texas. This climax forest range vegetation was conterminous with (immediately adjacent to) a Mexican pinyon pine (Pinus cerebroides)-alligator juniper (Juniperus deppeana)-Emory oak (Quercus emoryi)-mixed grass woodland (a savanna form of that range vegetation also). This latter sub-type or variant of pinyon pine-juniper woodland was included in the forest and woodland chapter of Range Types of North America entitled, Juniper-Pinyon Woodland.

52. About as many layers as imaginable- Physiogonomy, structure, and compostion of a ponderosa pine forest in the Sierra Blanca Mountain Range in southcentral New Mexico that had low shrub and herbaceous layers beneath widely speced pines of inetrmediate age. The main shrub--and, also, the associate species--was Gambel's oak (Quercus gambeli) while the second most abndant shrub was grey oak (Q. grisea). The herbaceous dominant was New Mexico sleepygrass (Stipa robusta). Other grasses included silver bluestem (Andropogon scoparius= Bothriochloa saccharoides= Bothriochloa laguroides ssp. torreyana), naturalized smooth brome (Bromus inermis), mountain or California brome (B. carinatus), fringed brome (B. cailiatus), and slender wheatgrass (Agropyron trachycauum). Forbs included purple sticky geranium (Geranium viscosissimum), giant milkvetch or giant locoweed (Astragalus giganteus), and meadow or blue flax (Linum lewisii).

This climax forest vegetation was the ponderosa pine/Gambel's oak (Pinus ponderosa/ Quercus gambeli) habitat type (Alexander et al, 1984, p. 14). It was a second-growth forest with trees of young adult age/size classes.

(On a separate ponderosa pine forest range, alligator juniper [Juniperus deppeana] and Fendler's muttongrass [Poa ffendleriana] were dominant understorey woody plant and herbaceous species, respectively. Those two important range plant species were also included below.)

53. Forest range par excellence- An open canopy, second-growth ponderosa pine forest range with an herbaceous understorey (foreground) and both an herbaceous and shrub layer (background) that developed in the Sierra Blanca Mountain Range in southcentral New Mexico. The dominant herbaceous species was New Mexico sleepy grass. Other grass species included silver bluestem; naturalized smooth brome along with mountain and fringed bromes, and slender wheatgrass. Forbs included purple, sticky geranium, giant milkvetch, and blue or meadow flax.Major shrubs were Gambel's oak with grey oak as the associate shrub species.

Portions of this ponderosa pine forest that did not have a woody component would be classified as the ponderosa pine-New Mexico sleepygrass (Pinus ponderosa-Stipa robusta) habitat type. In parts of this range vegetation where there was a shrub layer dominated by Gambel's oak it was a ponderosa pine/Gambel's oak (Pinus ponderosa/Quercus gambeli) habitat type (Alexander et al, 1984, p. 14).

54. All-in-one scenes- A second-growth ponderosa pine forest range with patches of a strictly herbaceaous understorey alternating with patches of both herbaceous and woody lower (shrubby) layers that developed in the Sierra Blanca Mountain Range in southcentral New Mexico. The dominant herbaceous species was New Mexico sleepygrass while the dominant shrub was Gambel's oak. The associate shrub was grey oak. Other herbaceous species included as the major grasses naturalized smooth bromegrass, mountain brome, fringed brome,and slender sheatgrss and such forbs as sticky geranium, giant milkvetch, and blue flax.

Overall, this forest range vegetation was a ponderosa pine/Gambel's oak (Pinus ponderosa/Quercus gambeli) habitat type (Alexander et al, 1984, p. 14). Patches devoid of a shrub component would be the ponderosa pine/New Mexico sleepygrass (Pinus ponderosa-Stipa robusta) habitat type.

55. Herbaceous and woody- Understorey of a second-growth ponderosa pine forest range in the Sierra Blanca Mountain Range in southcentral New Mexico with well-developed herbaceous and low woody layers. The overall dominant shrub was Gambel's oak while the dominant herbaceous species was New Mexico sleepygrass. In parts of this forest range that had a well-developed shrub layer (such as seen in these two images) the vegetation was a ponderosa pine/Gambel's oak (Pinus ponderosa/Quercus gambeli) habitat type (Alexander et al, 1984, p. 14). The associte shrub was grey oak.

Taxonomic note: the scrub oak now recognized as grey oak (Quercus grisea) was, previously, frequently known, grouped with, or, perhaps confused with, Q. undulata called by the common name, wavyleaf oak. Currently Q. undulata is recognized as a "catch-all name, without biological integrity" (Allred and IAvey, 2012, p. 350) for hybrids Q. gambelii and such species as Q. grisea, Q. turbinella, Q. muhlembergii, etc. (Allred and IAvey, 2012, p. 350; Carter, 2012, p. 313). Alexander et al. (1984, ps. 14-15) recognized a Pinus ponderosa/Quercus undulata habitat type.

56. Various shades- Shoots of a vigerous plant of New Mexico beardtongue (Penstemon neomexicanus) grew between equally vigerous plants of muttongrass or Fendler's bluegrass (Poa fendleriana) in the understory of a ponderosa pine forest range in the sierra Blance Mountain Range of southcentral New Mexico. This is in the Sacramento section of Basin and Range physiographic province. Exceptionally moist soil conditions were at least partly responsible for the large-sized, surperb specimens of both of the herbaceous range plants.

Muttongrass was the dominant species of the grassy understorey of this forest range while beardtongue was the major forb. The woody species closest to these herbaceous plants was alligator juniper (Juniperus deppeana) which was the most abundant midstorey plant species on this forest range.

57. Lucky to have anything- New Mexico sleepygrass (Stipa robusta) showed as a large specimen with shoots over six feet in height (first and faded-out slide; see shortly), basal shoots (second slide), and immature panicle inflorescence (third slide) growing in northcentral New Mexico

. Clearly (or unclearly was more like it) the first slide did not show much other than to give general scale of a mature plant (group of adult tillers) of New Mexico sleepygrass. The photographer's normal/macrolens started malfunctioning in the middle of what to him was a major photographic expedition. Hardened grease caused the aperature to stay open too long in various shots. The lens performed properly for the second and third slides in this series so there was still much to be thankful for.

Besides more photographs of Stipa robusta were presented below. These should "hold" viewers for the time being.

58. Muttongrass or Fendler's bluegrass (Poa fendleriana) is not only one of the Poa species having larger individual plants, but it also a major forage grass (perhaps related to its robust size) in various parts of the Western Range, including the interior ponderosa pine cover type. The following series of slides presented several plants (and reproductive parts of them) of muttongrass graowing in the understorey of a ponderosa pine range in the Sierra Blanca Mountain Range located in the Sacramento section of the Basin and Range physiographic province.

Good references for muttongrass included the timeless Range Plant Handbook (Forest Service, 1940, p. G100) as well as Tilley et al. (2006). By the way, the commemorative name for the specific epithet was for August Fendler who collected the type specimen (Wooton and Standley, 1915, p. 101; Forest Service 1949, p. G100)). Incidentially, there are probably more commemorative specific epithets based on surnames in Poa than in any other grass genus. (Just take a "gander" at any encyclopedia of North American grasses such as Hitchcock and Chase [1951] or Barkworth et al. [2007].)

The Range Plant Handbook remarked that muttongrass (and it has remarkable capacity to fatten sheep) was "one of the most widely distributed and important of native bluegrasses". The species range of muttongrass extends from British Columbia south to California and eastward to Nebraska and south to Texas and into northern Mexico (Hitchcock and Chase, 1951, p, 127). Muttongrass is clearly a climax (decreaser) species. It is generally dioecious ("incompletely dioecious" [Hitchcock and Chase, 1951, p. 126].

58a. For mutton (and just about everything else)- Several plants of muttongrass of Fendler's bluegrass, the dominant herbaceous species in the understorey of a ponderosa pine forest range in the Sierra Blanca Mountain Range. These comparatively large specimens were growing on a forest range from which livestock but not wildlife were excluded. There had been no grazing on these plants so they showed intact tillers (intravaginal or vertical shoots) most of which were sexual shoots with well-developed panicles loaded with grain.

These plants ranged in height from one and a half to two feet which was somewhat larger than typical for this species, but it had been a cool, moist spring following a wet winter so growing conditions were optimal for this cool-season, C3 festucoid grass species.

These (and all subsequent) slides were taken in late afternoon just before the sun starting "sinking down".

Lincoln National Forest, Lincoln County, New Mexico. Late June; peak standing crop and ripening caryopses.

58b. Ungrazed representatives- Some ungrazed plants of muttongrass or Fendler's bluegrass growing in the understorey of a pondrosa pine forest range from which livestock (and not wildlife) had been excluded. Almost all tillers were sexual shoots bearing large panicles with maturing grains (caryopses).

These unusually large speciments measured from one and a half to two feet in height. (Tilley et al. [2006] noted that muttongrass sometimes grows to a height of two and a half feet.) The specimens seen here were representative of muttongrass plants which dominated the herbaceous layer of a ponderosa pine forest in the Sierra Blanca Mountain Range located in southcentral New Mexico. Growing conditions, including soil moisture, had been extremely favorable (cool, moist spring following winter of heavy snow) for muttongrass such that these were larger-than-usual representatives.

Lincoln National Forest, Lincoln County, New Mexico. Late June; peak standing crop and ripening caryopses.

"Though nothing can bring back the hour
Of splendour in the grass, of glory in the flower;
We will grieve not, rather find
Strength in what remains behind..." --- William Wordsworth

58c. Splendor in grass- The above lines--perhaps reflecting Jesus' words as recorded in Matthew 6:30--have been frequently quoted and even served as the title for the picture show, Splendor in the Grass, but they were never any more literally true than in these examples of panicle-bering shoots of muttongrass of Fendler's bluegrass.

These ungrazed and unusually large specimens were growing in a ponderosa pine forest range on which this grass was the dominant herbaceous species of the understorey. This forest range was in the Sierra Blanca Mountains of southcentral New Mexico.

The amber-colored panicles reflected late afternoon sunlight just before onset of sunset.

Lincoln National Forest, Lincoln County, New Mexico. Late June; peak shoot development with ripening caryopses.

58d. Splendor up close- Parts of panicles (first slide) and spikelets in these panicles (second slide) of muttongrass or Fendler's bluegrass that was the dominant range plant in the herbaceous understorey of an interior ponderosa pine forest range in the Sierra Blanca Mountain Range located in southcentral New Mexico.

Fendler's muttongrass will frequently produce a heavy grain crop, but sometimes this is by apomixis (production of seed/fruit asexually--without fertilization--resulting in clonal daughter or sister offspring that in turn can become mother plants). Barkworth et al. (2007, ps. 556-559 passim) presented some details of sexual and asexual seed/grain production in the three subspecies of Poa fendleriana that they recognized and described.

Late afternoon light from a starting-to-set sun lite up the amber spikelets presented here.

Lincoln National Forest, Lincoln County, New Mexico. Late June; peak ripening caryopses.

59. New Mexico beardtongue (Penstemon neomexicanus) was the major or most abundant forb in the herbaceous understorey of a ponderosa pine forest range in the Sierra Blanca Mountain Range. Its closest neighbor was muttongrass or Fendler's bluegrass which was the dominant herbaceous speciesin the grassy understorey of this forest pasture from which livestock--though not wildlife--had been excluded. Allred and Ivey 2012, p. 443) stated that New Mexico beardtongue was endemic but not rare to mountains in this area of southcentral New Mexico.

It was a pleasnt find for your rangeman photographer who managed to collect the following photographs in late afternoon just before the sun sank too low for your textbook. "Quail and manna" from Heaven.

59a. Fading light and faded colors- Big (and almost rank-growing) plant of New Mexico beardtongue (Penstemon neomexicanus) in first slide and upper shoots of another plant in the second slide. These examples were growing beside muttongrass or Fendler's bluegrass in the herbaceous understorey of a ponderosa pine forest range in the Sierra Blanca Mountains in southcentral New Mexico.

Even in the rapidly fading light of early evening the color of the delightful pink corollas of this elegant range forb were faded out. Not to worry: these flowers and their pink pastel were captured and displayed in the very next set of two slides...

Lincoln National Forest, Lincoln County, New Mexico. Late June; mid-bloom phenological stage.

59b. No fading here- Flowers of New Mexico beardtongue which was the most abundant forb in the herbaceous understory of a ponderosa pine forest range in the Sierra Blanca Mountain Range in southcentral New Mexico.

Lincoln National Forest, Lincoln County, New Mexico. Late June; mid-bloom phenological stage.

60 a. A real geranium- A single plant (first slide) and upper shoot (second slide) of purple sticky geranium (Geranium viscosissimum) in the understorey of a second-growth ponderosa pine-Gambell's oak forest range in the Sierra Blanca Range in southcentral New Mexico. This range forb was one of three species in the herbaceous layer of this second-growth ponderosa pine forest. In North America the native Geranium species are almost never a major range plant species. One exception is forbland in some parts of the Rocky Mountain chain, especilly in the central Rockys. This range cover type was treated in its own chapter, Forbland, in this publication.

Even though Geranium species are not generally major range plants (in contrast to Erodium species, such as E. cicutarium,, which are also in the geranium family) they can be locally important as forage species and they do add species diversity to the range plant community beit forest, forbland, or ecotones.

It was Richardson's or wild white geranium (G. richardsonii) which the Society for Range Management included on its "200" (species) list for the International Collegiate Range Plant Identification Contest, but G. viscosissimum is very similar yet distinct with its pink to magenta flowers and usually sticky foliage (Hermann,1966, p. 170; Allred and Ivey, 2012, p. 359). Purple stick geranium was treated in Notes on Western Range Forbs in which Hermann (1966, p. 170) concluded that forage value of all the Geranium species was "highly variable".

The common domesticated "gernium" commonly seen in home flower gardens--usually as a bedding or potted plant is not a geranium at all but in the genus, Pelargonium which, however, is also in the geranium family (Geraniaceae). In fact, at one time Pelargonium species were included in the Geranium genus. In essence, the misnomer of "geranium" applied to the potted (and ill-scented) plant is not as far-fetched as might appear. Either way purple stick geranium is a true geranium and an understorey forest forb in the ponderosa pine-Gambel's oak range featured above.

Lincoln National Forest, Lincoln County, New Mexico. Late June; mid-bloom phenological stage.

60b. Gerany up- Upper shoot with flower of purple sticky geranium growing in the herbazeous layer or zone of a relatively open canopy, second-growth ponderosa pine-Gambel's oak forest range in the Sierra Blanca Mountain Range in southcentral New Mexico. This species was an occasional species on this forest range, but it certainly added biological diversity to the range--not to mention a beautiful color and, even, attractive (and distinctive) leaves.

No, purple stick geranium did not contribute any forage to speak of, but as Jesus reminded Satan in quoting from Deuteronomy 8:3, "man shall not live by bread alone" (Matthew 4:4, KJV). Rangemen learn to value the "priceless" aesthetics of the range as well as the practical, economic aspects (sometimes all a rangeman has is the aesthetic side).

Lincoln National Forest, Lincoln County, New Mexico. Late June; mid-bloom phenological stage.

60c. End-on and sideways- Views of the delightful magenta flower of purple sticky geranium from topdown view (first slide) and a side view (second slide). This flower was produced on the plant presented in the two immeidately preceding slide/caption units. It was "happily" blooming in the herbaceous layer of a second-growth ponderosa pine-Gambel's oak forest range in the Sierra Blacca Range in southcentral New Mexico (The Land of Enchantment of course.).

Lincoln National Forest, Lincoln County, New Mexico. Late June; mid-bloom phenological stage.

61.Now for a legume (and a damn big 'un)- Plant of giant milkvetch (Astragalus giganteus) growing in the herbaceous layer of a second-growth ponderosa pine-Gambel's oak forest range in the Sierra Blanca Range in southcentral New Mexico. This is a species of robust plants that have large woody taproots (Correll and Johnston, 1979, p. 845).

Giant milkvetch is NOT one of the known toxic Astragalus species (Burrows and Tyrl, 2013, 502-526). Rather it is a native, showy, nodulated (nitrogen-fixing), papilionaceous legume that, as suggested by the specific epithet, giganteus, is a species of characteristically large plants.

Lincoln National Forest, Lincoln County, New Mexico. Late June; early (maybe, only) bloom stage of phenology.

62. Big 'un in all regards- Papilionaceous Inflorescence of giant milkvetch. This is a beautiful Astragalus species that seemed to this author to show promish as an attractive species for use in landscaping with native plants. Even though this is a species of large forbs giant milkvetch has received comparatively little treatment compared to other Astragalus species. This is perhaps due to the fact that giant milkvetch is rather limited in its biological (species) range to a few mountain ranges in central New Mexico, a few areas in Trans Pecos Texas, and Chihuhua (Correll and Johnston, 1979, p. 845; Allred and Ivey, 2012, p. 311), and the fact that it is not a toxic Astragalus species (Burrows and Tyrl, 2013, 502-526).

Lincoln National Forest, Lincoln County, New Mexico. Late June; early (maybe, only) bloom stage of phenology.

63. Flax in the forest- Two examples (two individual plants) of blue or meadow flax (Linuum lewisii= L. perenne var. lewisii) growing in the understorey of a fairly open canopy, second-growth ponderosa pine-Gambel's oak forest range in the Sierra Blanca Mountains in southcentral New Mexico. In the herbaceous layer of which blue flax was a component species the dominant non-woody species was is New Mexico sleepygrass with native perennial Bromus species and silver bluestem.

Neither one of these plants showed evidence of feeding by any animal including insects except for visiting pollinators.

Lincoln National Forest, Lincoln County, New Mexico. Late June; mid-bloom stage of phenology.

64. Flexible flax- Upper shoots with flowers of blue flax . An interesting visual phenomenon was that color of petals varied not only from plant-to-plant and, even, on the same plant, but also of the same petals depending on light conditions. As indicated by one of the common names, blue flax, petals are typically light or pale blue to lavendar in color. The slightly windblown petals in the first slide were of a more common pale blue whereas the petals in the second slide were lavendar but with more of a pinkish hue when seen against a dark background and direct light hitting the corollas.

L. lewisii (obviously a commemorative name for Captain Merriwether Lewis of the famed Corps of Discovery, America's greatest scientific expedition) Blue or meadow flax is one of 15 Linuum species native to New Mexico (Allred and ivey, 2012, ps. 383-384)

Hermann (1966, p. 173) provided brief coverage of L. lewisii which has very little feed value under most conditions.

Lincoln National Forest, Lincoln County, New Mexico. Late June; mid-bloom stage of phenology.

Cuttin' loose on the floor- Cutleaf germander (Teucrium laciniatum) growing on the floor of a relatively open canopy, second-growth ponderosa pine-Gambel's oak forest range in the Sierra Blanca Mountains in southcentral New Mexico. This species joined other species in the herbaceous layer(s) including muttongrass, New Mexico sleepygrass, giant milkvetch, purple sticky geranium, blue flax, and purple beardtongue.

This low-growing, native member of the mint family (Labiatae) is widespread in the Land of Enchantment where it grows in varied habitats (Allred and Ivey, 2012, p. 380). Cutleaf germander was pretty common in this "neck of the woods".

Lincoln National Forest, Lincoln County, New Mexico. Late June; peak-bloom stage of phenology.

65. With and without sun- Two slides of two trunks of alligator juniper (Juniperus deppeana) rtaken with a matter of seconds of each other. The first shot (first slide) was in full light of a late afternoon sun (just before start of sunset) while the second shot (second slide) was right after a dense cloud passed between the subject and the sun. It is an age-old argument whether full-sun or overcast-sky shots produce "better" images. There is, of course, no simple answer for all situations.

This is a textbook example of that phenomenon. Probably most viewers would feel that in this specific instance the second slide (overcast-sky view) showed alligator juniper to better advantage, especially the needle foliage. On the other hand, plants of Fendler's muttongrass and three trunks of alligator juniper in the background were much better seen in the full-sun view. Your photographer author "compromised" and presented both views.

Lincoln National Forest, Lincoln County, New Mexico. Late June.

66. Seed and studded leaves- Fleshy seeds with their pointer protuberances and needles with their rough surfaces of alligator juniper in a ponderosa pine forest in which Fendler's bluegrass was the dominant of an herbaceous understorey. Alligator juniper grew as a lower tree or shrub layer in this relatively open forest range in the Sierra Blanca Mountain Range of southcentral New Mexico.

Lincoln National Forest, Lincoln County, New Mexico. Late June; mature seed stage.

Landscape at beginning of mid-elevation forest- A mid-elevation open ponderodsa pine forest with an understorey dominted by Gambel's oak in the Southern Rocky Mountain physiographic province. The first or upper slide presented the physiogonomy of the range forest vegetation and characteristic topography at its lowest point of contact with a stream and its narrow floodplain on which a willow carr (apparently of several Salix species) had developed.

The second or lower slide showed physiogonomy and overall structure of this forest range type just above the stream and its riparian plant community.

Jicarilla Apache Reservation, Rio Arriba County, New Mexico. Late July; mid-estival aspect. Forest range type was FRES No. 21 (Ponderosa Pine Ecosystem). Mapped by Kuchler as K- 17 (Ponderosa Pine-Douglas-fir Forest). SAF 237 (Interior Ponderosa Pine). No SRM for ponderosa pine cover types in the southwest region. Pinus ponderosa Association in Yellow Pine Series of Brown et al. (1998). Pinus ponderosa/Quercus gambellii habitat type (Ludwig et al., 1986). Southern Rockies- Sedimentary Mid-Elevation Forest ecoregion 21f (Omernik and Griffith, 2006). Willow carr and riparian zone of various Salix species in foreground of first or upper slide.

Movin' in on mid-elevation forest- The simple structure (basically three layers: 1) tree, 2) shrub, and 3) herbaceous) and general species composition of a ponderosa pine-Gambel's oak forest range plant community in the Southern Rocky Mountains in northcentral New Mexico. This and the next several sets of slides were of the Ponderosa Pine-Gambel's Oak habitat type (Ludwig et al., 1986).

This range had been heavily grazed--though not overgrazed/overbrowsed-- by elk (Cervus canadensis) and mule deer (Odocoileus hemionus), but influence (if any) on range vegetation was not known. Ponderosa pine and the scrub form of Gambel's oak obviously "overwhelmed" all other plants, except for local groups of comparatively young individuals of Douglas-fir (Pseudotsuga menziesii). In addition to these three conspicuous species there were a number of other range plant species. These included antelope bitterbrush (Purshia tridentata), true mountain-mahogany (Cercocarpus montanus), blue grama, galleta (Hilaria jamesii), little bluestem (Andropogon scoparius), tailcup lupine (Lupinus caudatus), cardinal beardtongue (Penstemon cardinalis), and various pre-bloom composites. There were also a few individual trees--and with very sparse cover at that--of Rocky Mountain juniper (Juniperus scopulorum) and alligator juniper (J. deppeana)

Jicarilla Apache Reservation, Rio Arriba County, New Mexico. Late July; mid-estival aspect. FRES No. 21 (Ponderosa Pine Ecosystem). Mapped by Kuchler as K- 17 (Ponderosa Pine-Douglas-fir Forest). SAF 237 (Interior Ponderosa Pine). No SRM for ponderosa pine cover types in the southwest region. Pinus ponderosa Association in Yellow Pine Series of Brown et al. (1998). Pinus ponderosa/Quercus gambellii habitat type (Ludwig et al., 1986). Southern Rockies- Sedimentary Mid-Elevation Forest ecoregion 21f (Omernik and Griffith, 2006).

Moved inside mid-elevation forest- Three views inside a Ponderosa pine-Gambel's oak habitat type forest range in the Southern Rocky Mountains of northcentral New Mexico. These three "photoplots" were of the same forest presented in the two immediately preceding slide sets. In these views there were very few Douglas-fir or juniper trees, but a sporadic herbaceous layer existed consisting of such species as blue grama, galleta, little bluestem, tailcup lupine, cardinal beardtongue (Penstemon cardinalis), and composites at pre-bloom phenological stages. There were some widely spaced plants of true mountain-mahogany and antelope bitterbrush.

Range vegetation had been heavily utilized by elk and mule deer, but not to he point of overbrowsing.

Tailed-up- Tail-cup lupine (Lupinus caudatus) growing in the understorey of a ponderosa pine-Gambell's oak forest in the Southern Rocky Mountains of northcentral New Mexico.

Jicarilla Apache Reservation, Rio Arriba County, New Mexico. Late July;peak-bloom phenological stage.

More inside views of mid-elevation forest- Two "photoplots" of the interior of a ponderosa pine-Gambel's oak (habitat type) game range heavily stocked with elk and mule deer though not to the point of overbrowsing. The herbaceous zone in forest vegetation here was mostly western wheatgrass (Agropyron smithii) with prairie sagewort (Artemisia frigida), western yarrow (Achillea lanulosa= A. millefolium), and redroot wild-buckwheat (Eriogonum racemosum) well-represented.

Views at outer margin of mid-elevation forest- Range vegetation on the outer edge of a ponderosa pine-Gambel's oak habitat type forest in the Southern Rocky Mountains in northcentral New Mexico. There plants of Rocky Mountain and alligator juniper along with antelope bitterbrush and true mountain mahogany. Locally major to dominant herbaceous species were western wheagrass, prairie sagewort, and redroot wild-buckwheat. Other herbaceous range plants included blue grama, galleta, pine dropseed (Blepharoneuron tricholepis), little bluestem, western yarrow, and tailcup lupine.

There was moderate to heavy utilization of browse plants by elk and mule deer, but no obvious evidence of overbrowsing.

An herbaceous patch- Two "photo quadrants" of an area of herbaceous vegetation in an open understorey ponderosa pine-Gambel's oak forest in the SouthernRocky Mountains in northcentral New Mexico. The two main species in this local spot wee western wheatgrass and prairie sagewort with western yarrow also common. Interestingly, there were also some small plants of the introduced pasture legume, bird's-foot trefoil (Lotus corniculatus).

There were seasonally high concentrations of mule deer and elk on this game range, but while browse utilization was high it was not at the point of overbrowsing. Herbaceous species could well have benefitted from the comparatively high degree of use on browse species including antelope bitterbrush and true mountain-mahogany as well as Gambel's oak.

Wild with red roots- Redroot wild-buckwheat (Eriogonum racemosum) growing in the understorey of a ponderosa pine-Gambel's oak forest with a high stocking rate of elk and mule deer in the Southern Rocky Mountains of northcentral New Mexico. Although not abundant so as to have much cover, plants of this range forb were widely scattered on this range as well as on ponderosa pine forests throughout this region.

Jicarilla Apache Reservation, Rio Arriba County, New Mexico. Late July; peak-bloom stage of phenology.

Deep inside an open forest- In the interior of a ponderosa pine-dominated forest with both lower woody (shrub) and herbceous layers that developed within the Southern Rocky Mountains in northcentral New Mexico. This highly structured forest (by ponderos pine forest standards) had Gambell's oak as the major (dominant) shrub, but wax or squaw current (Ribes cereum) was the associate shrub. (A well-developed patch of was current surrounding a ponderosa pine was presdented in center midground.)

Grasses included western wheatgrass, the dominan--often sole-- grass species along with Fendler's bluegrass or muttongrass, blue grama, galleta, and pine dropseed. Forbs were mostly prairie sagewort, western yarrow, and redroot wild-buckwheat.

Grass instead of shrubs- A open woodland-like form of ponderosa pine forest with a grass understorey co-dominated by Thurber's fescue (Festuca thurberi) and mountain muhly (Muhlenbergia montana) with Fendler's bluegrass or muttongrass, blue grama, sideoats grama, and pine dropseed comprising other important grasses. Local shrubs included Bigelow's sagebrush or flat sagebrush (Artemisia bigelovii) hairy grey gold-aster (Heterotheca villosa), and mountain big sagebrush (Artemisia tridentata subsp. or variety vaseyana). The main fobs were redroot wild-buckwheat and scarlet or red-flowere gilia, scarlet trumpet, or skyrocket (Gilia aggegata= Ipomopsis aggregata).

Carson National Forest, Rio Arriba County, New Mexico. Late July; peak standing crop and grain-shatter stages of phenology in grass species.

NOTE: slides of a ponderosa pine-grass understorey in Carson National Forest were taken in late evening and with a cloudy overcast. Hence, poor quality photographs, but no alternative except no slides at all.

A grassy underneath and glades- Open areas within an already sparsely tree-covered ponderosa pine forest in the Southern Rocky Mountains of northern new Mexico was co-dominated by mountain muhly and Thurber's fescue which are both bunchgrasses. Other grass species included Fendler's bluegrass or muttongrass, blue grama, sideoats grama, and pine dropseed. comprising other important grasses. Sparse forb cover was limited primarily to redroot wild-buckwheat and scarlet or red-flowere gilia. Widely scattered shrub species were mostly small plants of Bigelow's sagebrush or flat sagebrush and hairy grey gold-aster with a few larger individuals of mountain big sagebrush.

Carson National Forest, Rio Arriba County, New Mexico. Late July; peak standing crop and grain-shatter stages of phenology in grass species.

Details of a grassy underneath- Herbaceous layer and forest openings (= glades) in a ponderosa pine forest range in Southern Rocky Mountains. Co-dominant herbaceous species were Thurber's fescue and mountain muhly with Fendler's bluegrass or muttongrass, blue grama, sideoats grama, and pine dropseed rounding out the grass component. Almost all of these range plant species were discernable in the first "photo-quadrant" with a plant of mountain big sagebrush conspicuous in left center.

The three cespitose plantsin foreground of the second slide were Thurber's fescue. A plant of scarlet trumpet, red-flowere gilia, or skyrocket and several plants of Bigelow's sagebrush were also visible in this second slide.

Carson National Forest, Rio Arriba County, New Mexico. Late July; peak standing crop and grain-shatter stages of phenology in grass species.

Another namesake for Thurber- Four cespitose individuals or specimens of Thurber's fescue (Festuca thurberi) growing in the grassy understorey of an open ponderosa pine forest range whee it was co-dominant with mountain muhly. Barkworth et al. (2007, p. 408-411 passim) described Thurber's fescue as a big, "densely cespitose" bunchgrass.

Thurber fescue was regarded as a valuable range plant, especially for forage (and more so for cattle and horses than for small ruminants), in the Range Plant Handbook (Forest Service, 1940, G64). Allred and Ivey (2012, ps. 662-663) remarked that Thurber's fescue provided good forage for livestock though it was less plentiful than Arizona fescue (Festuca arizonica). Both are grasses of higher (subalpine) meadows and montane forests.

Thurber's fescue has been recognized in New Mexico since the earliest botanical (floral) treatments (Wooten and Standley, 1915, p. 102). It was included with (or shown synonymous with) rough fescue (F. scabrella) in the Rocky Mountain manual by Colter and Nelson (1909, p. 76).

George Thurber served as botanist, quartermaster and commissary on on the famed United States and Mexico international border expedition known as the United States and Mexican Boundary Survey West (1848–1855) following signing of the Treaty of Guadalupe Hidalgo in 1848.

Carson National Forest, Rio Arriba County, New Mexico. Late July; peak standing crop and early grain-shatter stage of phenology.

Thurbered tops- Panicles of Thurber's fescue that were produced in the herbaceous understorey of a ponderosa pine forest in the Southern Rocky Mountains of northern New Mexico.

Carson National Forest, Rio Arriba County, New Mexico. Late July; peak standing crop and early grain-shatter stage of phenology.

Grass of low mountains- Mountain muhly (Muhlenbergia montana) in an open ponderosa pine forest with a grassy understorey co-dominated by Thurber's fescue and mountain muhly. The first of these three slides presented a large, characteristic plant of this primarily tufted (= cspitose) species. Barkworth et al. (2003, ps. 183-184) stated that mountain muhly was not rhizomatous, but the Forest Service (1940, G81) stated that mountain muhly does have short rhizomes (Forest Service (1940, G81) so it it is not strictly cspitose or as pronounced a bunchgrass as some species. This author would put his money on the Forest Service (1940, G81) over Barkworth et al. (2003) almost any time. Presence of rhizomes was indicated in the first (and to lesser extent, in the second) image in which smaller bunches of daughter tillers were visible arising from short rhizomes.

The third (bottom) slide in this trio of images showed the sparsely branched, spreading panicle type inflorescence with long, curveed awns at grain-ripe are readily identifiable features of this valuable climax forage grass.

As noted above mountain muhly was discussed as a range plant in the Range Plant Handbook (Forest Service, 1940, G81). This importance of mountain mhly was reflected by its inclusion as one of the 200 species on the International Rane Plant Identification Contest sponsored by the Society for Range Management Stubbendieck et al., (1992, p. 124-125). Mountain muhly (at least not as M. montana) was not referenced in the earliest floral treatise of New Mexico (Wooten and Standley, 1915) nor in the Rocky Mountain manual of Colter and Nelson (1909).

These slides were of necessity taken near sundown so that photographic quality is far from superb, but it was what a rangeman so blessed to experience would see. Rangemen can regard this sunset light as the shining face of Zia, the sun symbol of the Zia Pueblo tribe. The Zia symbol of this southwestrn Indian tribe became the adorning symbol on the New Mexico state flag. To the Zia people their symbol signified what they regarded as the great Circle of Life with the four seasons, four geographic directions, and four periods of each day. These sun-glare images reminded students that energy from the sun as light is the source of all life on the range. It is notable that so many of man's religions regard light as symbolic or the manisfestation of God, the Supreme Being, by whatever name.

Carson National Forest, Rio Arriba County, New Mexico. Late July; peak standing crop and grain-shatter phenological stage.

A muhly panicle- Panicle of mountain muhly taken from one of the plants presented immediately above. The spreading inflorescence was laid on part of a shattered limb of ponderosa pine in this open Rocky Mountain ponderosa pine forest range.

Carson National Forest, Rio Arriba County, New Mexico. Late July; peak standing crop and early grain-shatter stage of phenology.

Transition forest- Lower elevation montane forest at the upper elevational limit of Douglas-fir and lower elevational zone of ponderosa pine in Sangre de Cristo Mountains, part of a forest range at the southrnmost end of the Southern Rocky Mountains. Shrubs in these two photographic forest profile views were true mountain-mahogany (Cercocarpus montanus). There were open grassy patches in the understorey of this forest (as shown in the immediately following slides).

Carson National Forest, Taos County, New Mexico. Late July; mid-estival aspect. FRES No. 21 (Ponderosa Pine Ecosystem). Mapped by Kuchler as K- 17 (Ponderosa Pine-Douglas-fir Forest). SAF 237 (Interior Ponderosa Pine). No SRM for ponderosa pine cover types in the southwest region. Pinus ponderosa Association in Yellow Pine Series of Brown et al. (1998). Pinus ponderosa/ habitat type (Ludwig et al., 1986). Southern Rockies- Sedimentary Mid-Elevation Forest ecoregion 21f (Omernik and Griffith, 2006).

Valuable dominant shrub- Shoot, leader with leaves, and fruit (first, second, and third slide, respectively) of true or common mountain-mahogany (Cercocarpus montanus), the local dominant shrub in a ponderosa pine forest range in the Sangre de Cristo Mountains in northern New Mexico. The Cercocarpus species, especially C. montanus, are some of the most valuable browse plants in North America, especially on ranges of the western part of the continent. Carter (2012, p. 417) provided a recent treatment of true or common mountain-mahogany.

True mountain-mahogany is one of 200 range plant species included on the Range Plant Identification Contest sponsored by the Society for Range Management (Stubbendieck et al., 1992, ps. 406-407).

Carson National Forest, Taos County, New Mexico. Late July; mid-maturity fruit phenological stage.

Great Plains-Rocky Mountains Super Combo-Young adults of ponderosa pine and saplings of Dougls-fir with an understorey of western wheatgrass comprised a forest in the Southern Rocky Mountains at the edge of the tablelands portion of the Southern Great Plains (Llano Estacado or High Plains). Ponderosa pine is the climax dominant of mid-elevation Rocky Mountain forests while the mid-grass, western wheatgrass, is one of the regional dominants of the Mied Prairie.

The highly rhizomatous western wheatgrass had formed patches of essentially single-species stands. In the well-lite interior of this open forest Douglas-fir saplings grew in roundish versus the triangular, spire-shaped form that is more typical of this fairly shade intolerant conifer.

The tallgrass species were Texas bluestem (Andropogon cirratus= Schizachyrium cirratum) and bull muhly or bullgrass (Muhlenbergia emersleyi). Major midgrasses were cane bluestem (Andropogon barbinodis= Bothyriochloa barbinodis), sideoats grama (Bouteloua curtipendula), green sprangletop (Leptochloa dubia), plains bristlegrass (Setaria leucopila), and plains lovegrass (Eragrostis intermedia). , Blue grama (B. gracilis) and pinyon ricegrass (Piptochaetium fimbriatum) were the only shortgrass species, but their presence gave a tallgrass, midgrass, and shortgrass component to the herbaceous understorey so as to make this "grass zone" a "complete mixed prairie". The overall dominant grass was cane bluestem, but others such as Texas bluestem and bullgrass were local dominants.

Forbs were limited, but big plants of giant goldenrod (Solidago gigantea) grew impressively along margins of a small stream tht drained this forest range..

This ponderosa pine forest range had beeen beset by several years of drought (Palmer Drought Severity Index in the Severe Drought to Extreme Drought stages) after which the native coleopteran, western pine beetles (Dendroctonus brevicomis= D. ponderosa) invaded the forest and killed from 40% up to 60% of the ponderosa pines. Then a wildfire swept through this forest of of readily flammable material (drought-killed herbage and standing, beetle-killed pines) to take out still more ponderosa pines. Officials with the Texas Forest Service (personal communication) estimated that over 90% of all ponderosa pines had been killed by this combination of stress factors. Since this "three-prong attack" there had been very little regeneration of pine. There is going to be human-replanting of ponderosa pine seedlings produced from local seed sources, but it will be decades before this dominant climax tree species recovers on one of the most xeric habitats on which it can survive. An extremely harsh forest range site, but one wonders who (what) replanted the pines killed by this same combination of natural catastrophes in the prehistoric past.

The western pine beetle is a native species that has had ecological effects on pine- and other conifer-dominated forests of western North America for countless millions (at least tens of thousands of years) as explained in numerous reviews such as those ranging from Shelford (1963, ps. 179-180) through Stark and Dabistento (1970) to DeMars and Roettgering (1982). This did not, of course, make the ponderosa pines on this devestated forest range any less dead. It did, however, illustrate in about the most dramatic example possible how the ravages by Mother Nature influence naturalforest vegetation.

Meanwhile the native grasses on this range were having a proverbial "field day" following one of the heaviest rainfalls ever recorded for the warm-growing season.

U up-U down Ranch built by the G. S. Locke family, now Davis Mountains Preserve of The Nature Conservancy, Jeff Davis County, Texas. Early October, early autumnal aspect. FRES No. 21 (Ponderosa Pine), K-Western Ponderosa Pine), SAF 237 (Interior Ponderosa Pine); no SRM for ponderosa pine cover type(s) in the Southwest. Pinus ponderosa Association in Yellow Pine Series of Brown et al. (1998). Arizona/New Mexico Mountains- Montane Woodlands Ecoregion, 23b (Griffith et al., 2004).

68. Multi-layered savannah form- A ponderosa pine-dominated forest range cover type with pines of such low density or widely spaced dispersion that it was more of a savanna form (certainly no more than woodland structure). Alligator juniper and a few redseed juniper along with a "respectable" cover and density of Emory oak completed the list of tree species on this forest range. Aside from shrub-sized and shrub-form of Emory oak and of the two juniper species, the shrub component was completed by Texas madrone. Grass species, beginning with the dominant cane bluestem, included sideoats grama, plains lovegrass, green sprangletop, blue grama, wolftail, Texas bluestem and, on moist banks of a small stream, bullgrass or bull muhly. Bullgrass was joined by large plants of giant goldenrod, about the only forb of much consequence on this forest range.

The more orange color of the understory as seen in the second of these two slides was the herbage of Texas bluestem (see four slide/caption sets below). Most of the green crowns in these two slides was that of Emory oak with lesser cover of alligator juniper.

Foresters with the Texas Forest Service (personal communiction) estimated that at least 90% of the ponderosa pines had been killed by a combination of drought, wildfire, and western pine beetles. Restoration efforts were underway to plant ponderosa pine seedlings from local germ plasm of this dominant and defining native plant of this climax plant community.

69. A few ponderosa pines left after onslaught by three foes- A woodland (or more like a savanna) form of a climax ponderosa pine-mixed grass range community in the Davis Mountains of Trans-Pecos Texas as it appeared after a multi-year drought ranging from Severe to Extreme (Palmer Scale), two recent wildfires, and attacks by western pine beetles. About 90% of the ponderosa pine on this forest range had died as a combination of these three compounding disturbances. The green tree and shrub crowns seen in these two images were those of Emory oak and, with lesser cover, alligator juniper.

The herbaceous understorey presented in thee two slides was exclusively that of native, perennial, midgrass species, the dominant of which was cane bluestem. Associate grass species varied at local scale and included sideoats grama, green sprangletop, plains lovegrass, and Texas bluestem. There was a small proportion of herbaceous cover in the shortgrass species of blue grama, pinyon ricegrass, and wolftail (this latter could be regarded as a midgrass).

This forest range had served as natural pasture for beef cattle for over a century, but it had been destocked of cattle for several years, including at time of these photographs. Both white-tailed and mule deer were free-ranging on this savanna form of a climax ponderosa pine-alligator juniper-Emory oak-mixed grass forest.

This forest range had been destocked of beef cattle for the past several years, but it had served as range for beef cattle for a century or more. The climax range plant community shown here had come through a multi-year drought (Severe to Extreme Drought on the Palmer Scale) and two wildfires over the last five years. Not bad vegetative recovery, hugh! Obviously riparian environments would be expected to recovery from disturbances faster than adjoining, less mesic habitats, but "nieghboring" slides in this section attest to the amazing ability of native range plants to recover quickly on less mesic habitats once favorable growing conditions return--as they always do.

This view, like all those above and below it, was taken at end of one of the wettest summers in weather records.

71. Dead trees; thriving grass- On a climax ponderosa pine-alligator juniper-Emory oak-mixed grass forest in the Davis Mountains of Trans-Pecos section of the Basin and Range physiographic province a powerful combination of multi-year drought (Severe to Extreme on the Palmer Index), two wildfires, and attacks on ponderosa pine by western pine beetle left less than 10% of ponderosa pine present prior to this three-factor disturbnce phenomenon based on estimates by foresters of the Texas Forest Service (persona communication). Even some alligator juniper had been killed outright or at least topkilled by wildfires. Emory oak survived.

As for the native, perennial, warm-season grasses? They not only survived, but appeared to be thriving at end of one of the wettest summers in weather records. The featured grass plant in this scene was bullgrass or bull muhly (center foreground) growing beside the crashed crown of a ponderosa pine that was not so lucky (not so well-adapted gtenetically, perhaps). Other grass species present included cane bluestem (the overall dominant herbaceous species), Texas bluestem, green sprangletop, sideoats grama, and plains lovegrass. Shortgrass species like blue grama and pinyon ricegrass were not visible in the luxuriant stand of midgrasses seen here.

72. Cane under ponderosa pine- Cane bluestem (Andropogon barbinodis= Bothriochloa barbinodis= B. b. var. barbinodis, A. torreyanus), local dominant to co-dominant grass in the understorey of a ponderosa pine woodland or savanna in the Davis Mountains of Trans-Pecos Texas. These plants (and those featured in the immediately following two slides) had grown their annual shoots in a rainfall record-setting warm-growing season following several years of Severe to Extreme Drought.

Cane bluestem is the major bluestem--and, often, the dominant, grass species--growing in the mountains of the arid zone of southwest North America. Powell (2000, p. 329) stated that cane bluestem "is perhaps the most widely distributed bluestem in the Trans-Pecos". Cane bluestem is the ecological dquivalent of the very similar silver bluestem (A. saccharoides= B. saccharoides= B. laguroides) which is co-dominant with sideoats grama over much of the Southern Great Plains mixed prairie. Cane bluestem (as A. saccharoides) has been recognized for this region since publication of Botany of Western Texas (Coulter 1891-1894, p. 497), the earliest flora or manual of this vast territory, and the also classic Flora of New Mexico (Wooton and Standley, 1915, p. 50).

Cane bluestem or cane beardgrass was also recognized (by one of the above species names) and dscribed by the following agrostologists: Silveus (1933, p.716-717), Hitchcock and Chase (1951, p. 768, 769), Gould (1975, p. 600-601), Powell (2000, p. 328-329), Barkworth et al. (2003, ps. 642-644 passim), Allred and Ivey (2012, ps 638, 640), and Shaw (2012, p. 257). Cane bluestem is also a major bluestem species westward to the Pacific Ocean in southern California. It is a major grass species in the general region of the Sonoran Desert and adjacent foothills and mountains (Kearney and Peebles, 1960, p. 142; Shreve and Wiggins, 1964, p. 298-299). Cane bluestem was recognized as an important range plant in the Range Plant Handbook (U.S. Forest Service, 1940, G14).

There are a number of Andropogon species in section 3 Amphilophis of Hitchcock and Chase (1951, p. 751), plus several additions of introduced species since that time, that are extremely similar to each other and have been split out (now as separate Bothriochloa species) based only on minute structural details. For example, Shaw (2012, p. 255) separated all of these morphologically similar Bothriochloa species in Texas based on sessile spikelets less than or more than 4.5mm in length and within these two dichotomies rachis lengths, leaf blade widths, and pubescence features.

Cane bluestem has been rated as being of Fair to Good forage value (in general). For Arizona ranges Humphrey (1960, p. 9) recommended grazing cane bluestem during active summer growth when palatability and nutritive content were greater. Humphrey (1960, p. 9) also recommended leaving a third of the sexual shoots of cane bluestem to provide seed for new plants as well as for maintenance of vigerous root systems.

U up-U down Ranch built by the G. S. Locke family, now Davis Mountains Preserve of The Nature Conservancy, Jeff Davis County, Texas. Early October; peak standing crop.

73. Featuring shoots of cane (bluestem, that is)- Shoots of cane bluestem growing in the understorey of a ponderosa pine savannah or woodland in the Davis Mountains of southwest Texas, part of the immense Basin and Range physiographic province. The remarkable size and quantity of herbage produced here took place in a record-setting rainfall period that extended over the entire warm-growing season. The only larger herbivore species that grazied this forest (savannah) range were white-tailed deer (Odocoileus virginianus) and mule deer (O. hemionus).These deer probably consumed very little of this herbage, but the specimens presented here and in the immediately preceding slide demonstrated the amount of potential forage that can be produced in a "good year" by cane bluestem on ponderosa pine forest ranges.

Forage value and proper management for cane bluestem was given in the preceding caption.

U up-U down Ranch built by the G. S. Locke family, now Davis Mountains Preserve of The Nature Conservancy, Jeff Davis County, Texas. Early October; peak standing crop.

74. Dead give-away; dead ringer- General view of portions of several tillers of cane bluestem that grew in the herbaceous understorey of a ponderosa pine savanna range in the Davis Mountains of Trans-Pecos Texas. These shoots had been produced in a record-shattering wet warm-growing season. Even at this distance the distinctive circles of white hairs encircling the culm of this species were visible. This ciliate (probably hairs not long enough to be described as villous) pubescence is a "dead-give-away" feature of this species. This telltale feature was presented at closer camera-range in the next two slides …

U up-U down Ranch built by the G. S. Locke family, now Davis Mountains Preserve of The Nature Conservancy, Jeff Davis County, Texas. Early October; peak standing crop.

75. Dead ringer details- Phytomers (the node-internode units) of cane bluestem tillers showing the distinctive (and sort of cute) ring of ciliate pubescence that grew around and completely encircled the culm of cane bluestem. These shoot units were on the same tillers as presented in the immediately preceding slide. They were on plants growing in the herbaceous understorey of a ponderosa pine savanna (woodland, perhaps) in the Davis Mountains of Trans-Pecos Texas.

U up-U down Ranch built by the G. S. Locke family, now Davis Mountains Preserve of The Nature Conservancy, Jeff Davis County, Texas. Early October; peak standing crop.

76. Happy and thriving cane tops- Two views of the inflorescence (numerous closely spaced rames) of cane bluestem (with a few shoots of green sprangletop [Leptochloa dubia] for a sidekick) in the Chihuhuan Desert scrubland. The inflorescence (flower cluster) of cane bluestem has more recently been described as a panicle of racemose primary and, sometimes, secondary branches (Powell, 1988, p. 324). This arrangement has, in the perspective of some agrostologists, qualified bluestem species having this floral arrangement to be in the genus Bothriochloa rather than the traditional Andropogon.

These shots from the adjoining Chihuhuan Desert showed the adaptation of cane bluestem as a climax (decreaser) dominant species in the Trans-Pecos Region. It was explained above that cane bluestem is also a dominant grass species in parts of the Sonoran Desert and adjoining foothills and forest ranges. The biological (species) range of cane bluestem extends westward to the Pacific Ocean (Kearney and Peebles, 1960, p. 142; Shreve and Wiggins, 1964, p. 298-299; Barkworth et al., p. ).

Big Bend National Park, Brewster, County, Texas. Early October; early autumnal aspect in a very wet year.

77. Lone Star Bluestem- Texas bluestem (Andropogon cirratus= Schizachyrium cirratum) in God's Country. Texas bluestem (as A. cirratus) was recognized for the Trans-pecos Region early on by Coulter (1891-1894, p. 495) and (as both A. cirratus and S. cirratum) by Wooton and Standley (1915, p. 50). Other authors who recognized this unique species included Silveus (1933, p. 715-716), Hitchcock and Chase (1951, p. 753), Gould (1975, p. 606), Powell, (2000, p. 232), Barkworth et al. (2003, ps. 674, 676, 677), Allred and Ivey (2012, p. 684), and Shaw (2012, p. 868). In their arrangement of Andropogon by sections Hitchcock and Chase (1951, p. 749) included A. cirratus in section 1. Schizachyrium. Since then most authorities treated Texas bluestem as Schizachyrium cirratum. "Same difference" as it were.

In addition to the above agrostologists, cane bluestem was recognized as an important bluestem in the greater Sonoran Desert Region, including foothills and mountain forest ranges as well as mostly upper elevations of the Sonoran Desert itself (Gould, 1950, 307-308; Kearney and Peebles, 1960, p. 142; Shreve and Wiggins, 1964, p. 299). With plants capable of acheiving sizes seen here (and in the immediately following two slides) Texas bluestem would have to be considered as having Good forage value, at least until proven otherwise (there seem to have no animal diet or utilization studies that examined palatability of Texas bluestem), but Humphrey (1960, p. 10) explained that due to "its rather low palatability" Texas bluestem "usually rates as only fair forage, probably because it generally grows among highly palatable grama grases". For Arizona ranges, Humphrey (1960, p. 10) recommended grazing in the summer months of July through September while leaving one/third of the sexual shoots for grain production and maintenance of good root systems.

Under the right conditions most of the bluestems (Andropogon species, including all the post-Hitchcock sectional split outs) turn beautiful colors in late summer and autumn, but Texas bluestem is in a limited league even among these. This photographer cannot recall ever seeing any more spectacular autumn- colored grass--any place at any time--than specimens of Texas bluestem in the Davis Mountains of Trans-Pecos Texas which is the eastern side of the huge Basin and Range physiographic province.

If you think this plant was beautiful just advance the carousel to the next two slides…

U up-U down Ranch built by the G. S. Locke family, now Davis Mountains Preserve of The Nature Conservancy, Jeff Davis County, Texas. Early October; peak standing crop, post grain-shatter phenological stage.

78. Local understorey dominant and its sidekicks- Texas bluestem, a local herbaceous dominant, with three associated grasses--plains bristlegrass (Setaria leucopila), sideoats grama (Bouteloua curtipendula), and blue grama (B. gracilis)--in the understorey of a ponderosa pine-dominated open canopy forest (or, more accurately, a savanna). This climax woodland (savanna) vegetation was in the Davis Mountains of the Trans-Pecos portion of the Basin and Range physiographic province.

This range had not been grazed by livestock in several years, but white-tailed deer and mule deer ran freely on it. Prior to that time beef cattle had grazed this range for at least a century. A prolonged multi-year drought (varied from Severe to Extreme Drought rating on the Palmer Index) combined with wildfire had taken place over course of the immediate past several years except for the current warm-growing season for which there had been near-record rainfall. Range plants have evolved so as to respond rapidly once favorable growing conditions return following stress-filled periods.

As promished immediately above, this was some of the most spectacularly beautiful autumn foliage--anyplace anywhere. Texas bluestem is one of the most colorful--and just plain beautiful--of all the bluestems all of which can have shoots of brillant autumn coloration.

U up-U down Ranch built by the G. S. Locke family, now Davis Mountains Preserve of The Nature Conservancy, Jeff Davis County, Texas. Early October; early autumnal aspect, peak standing crop, post grain-shatter phenological stage. FRES No. 21 (Ponderosa Pine), K-Western Ponderosa Pine), SAF 237 (Interior Ponderosa Pine); no SRM for ponderosa pine cover type(s) in the Southwest. Pinus ponderosa Association in Yellow Pine Series of Brown et al. (1998). Arizona/New Mexico Mountains- Montane Woodlands Ecoregion, 23b (Griffith et al., 2004).

79. Regrouping after fire and drought- Resprout shoots from trunk base or rootcrown of a Texas madrone on a climax ponderosa pine-alligator juniper-Emory oak-mixed grass forest in the Davis Mountains of Trans-Pecos Texas section of the Basin and Range physiographic province after two wildfires and on-goining, prolonged drought for three and four years prior to time of this photograph. In other words, this was two years of regowth of Texas madrone. In the second year of that regrowth, which was the current year of this slide, rainfall during summer had been near-record amounts. The previous year had been the end of a multi-year drought that was rated as Severe to Extreme (Palmer Index). There was also drought when the wildfires occurred.

Over 90% of the ponderosa pines growing on this range were killed by the combination of drought, western pine beetle attack, and wildfire (foresters, Texas Forest Service, personal communication).

Surrounding climax grass included cane bluestem, sideoats grama, green sprangletop, plains lovegrass, Texas bluestem, blue grama, pinyon ricegrass, and wolftail.

80. Front Range of Colorado Rockies— This landscape scale view of the Montane life zone on the east side of the Continental Divide shows ponderosa pine going form open forms that vary from savanna to woodland to closed canopy forest. There are also open meadows of both Great Plains and mountain grasses plus Douglas fir and lodgepole pine (Pinus contorta) forests. Elevation 8,500 feet. Rocky Mountain National Park, Larimer County, Colorado. August. 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. Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

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.

81. 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).

82. 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).

83. Blooms on the Front Range- Inflorescences of thinbleberry on the plant presented in the preceding slide.Rocky Mountain National Park, Larimer County, Colorado. Mid-June, full-bloom stage.

84. 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.

85. 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.

86. 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."

87. 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.

88. 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?

89. Park-like physiogonomy of ponderosa pine montane forest with both shrub and herb layers. Mountain mahogany and wax current (Ribes cereum) are major shrubs. Herbs are primarily grasses of the Great Plains like blue grama, Junegrass, and western wheatgrass with some forest species like mountain brome (Bromus marginatus) and timber oatgrass (Danthonia intermedia). Larimer County, Colorado. August. FRES No. 21 (Ponderosa Pine Ecosystem). K-17 (Ponderosa Pine- Douglas-fir Forest). SAF 237 (Interior Ponderosa Pine); no SRM. Pinus ponderosa Association in Yellow Pine Series of Brown et al. (1998). Southern Rockies- Crystalline Mid-Elevation Forests Ecoregion, 21c (Chapman et al., 2006).

90. 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).

91. 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).

92. 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.

93. 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).

94. 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).

95. 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).

96. 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).

97. 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.

98. 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.

99. 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.

100. 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.

101. 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).

102. 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.

103. 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).

104. 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).

105. 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).

106. 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.

107. 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.

108. 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.

109. 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.

110. 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.

111. 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

112. 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.

113. 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).

114. Young specimens-Two examples of immature spikes in slender or bearded wheatgrass.

Pike National Forest, Teller Counter, Colorado. Early July; milk stage of grain development.

115. 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.

116. 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.

117. 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.

118. 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).

119. 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.

120. 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.

121. 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.

122. 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.

123. 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.

124. 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).

Taxonomic note: recent phylogenetic arrangement (Barker et al., 2007, 114-142, 156-184 passim) based on cladistics placed Indian ricegrass in the same genus as that of subalpine needlegrass and sleepygrass, a taxonomic treatment at considerable odds with traditional systematic interpretations and one which this author elected to disagree and ignore.

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

126. 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.

127. 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.

128. 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.

129. 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.

130. 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.

131. 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.

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

132. 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).

133. 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.

134. 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.

135. 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.

136. 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.

137. 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.

138. 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.

139. 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 County, 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).

140. 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.

Pike National Forest, Teller County, 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).

141. 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).

142. 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 County, 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).

143. 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 County, Colorado. Early July; mature fruit just prior to shedding.

144. 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 County, Colorado. Early July; mature fruit just prior to shedding.

Ponderosa Pine Forests and Adjoining Riparian Vegetation

Forests dominanted by Douglas-fir (Pseudotsuga menziesii) typically develop at lower to middle elevations on more mesic environments (= forest range sites) such as bottomlands or floodplains and north slopes.This phenomenon exist for both Interior Douglas-fir forest cover type (SAF 210) and the Pacific Douglas-fir forest cover type (SAF 229) (Eyre, 1980).

The following section dealt with Interior Douglas-fir in the Southern Rocky Mountains. Given development of Douglas-fir forests on bottomlands these mesic-habitat forests are often in direct contact (contiguous or conterminous with) riparian zones such as those comprised primarily of willow (Salix) species. In fact, Douglas-fir may itself be a riparian tree species and, hence, a component of riparian communities (eg. an associate to the dominant Salix species).

A tale of two types- A riparian shrub community made up of red or Pacific willow (Salix lasiandra= S. lucida subsp. lasiandra), thinleaf or mountain alder (Alnus tenuifolia= A. incana subsp. tenuifolia), water birch (Betula occidentalis), and Rocky Mountain maple (Acer glabrum), plus some herbaceous species including cutleaf coneflower (Rudbeckia laciniata) and sleepygrass (Stipa robusta) fronted a second-growth, north slope Douglas-fir forest along a stream in the Sangre de Cristo (Blood of Christ) Range. The Sangre de Cristo Mountains are the southermost extension of the Rocky Mountains. The Sangre de Cristo range or subrange is in southern Colorado and northern New Mexico.

Douglas-fir dominated forest are often some of the lowest elevation forest range types across much of western North America. Habitat of Douglas-fir forests includes bottomlands, often being part of the greater riparian zone vegetation, and north-slopes which, as in the example seen here, are contiguous with streamside plant communities.

Grazing value of Douglas-fir-dominated forest vary considerably ranging from high quality understorey range (often of both lower woody and herbaceous layers) where there is lower stocking density and less crown cover of trees (Douglas-firs are more are more widely spaced) to a non-existent range resource closed forest canopy cover and very limited light penetrating to the understorey. The latter condition is more apt to prevail in forests of older, larger trees (as in old-growth stands). In such old-growth Douglas-fir forests, or even stands at advanced successional stages with mature trees, the native grazing land is transitiory forest range (ie. range is in transition to closed forest canopy so that forest range exist only under certain, less advanced seral stages). This is in contrast to permanent forest range where a grazable/browseable understory--perhaps of several vegetational layers--persist throughout the entire successional sequence through to climax vegetation.

In contrast to Douglas-fir forest range (even the less common permanent forest range), range afforded by riparian vegetation almost always supplies relatively high-quality feed (browse and/or herbage), and it is usually permanent not transitory range). Given 1) the high-quality diets afforded by riparian vegetation, 2) inherent "laziness" of range animals (propensity to hand in on the more level, lower land along streams), and 3) general lack of predators (at least in sizeable populations) riparian plant communities have historically been overbrowsed/overgrazed and become sacrifice range sites.

Much of this damage has been reduced or, even, eliminated, especially on public land where resource-users, besides fee-paying stockmen, treasure streams and their protecting vegetation for recreation including sport fishing. For their part, stockgrowers responded to the challenge of stream degradation--even on private property. Provision by livestock associations of high-profile, prestiguous stewartship awards for proper livestock mangement has, in many instances, fostered dramatic recovery of stream habitats due mostly to restoration of riparian vegetation. People respond to incentives, be these internal or external.

Santa Fe National Forest, Taos County, New Mexico. Late July; mid-estival aspect. Forest cover type: FRES No. 21 (Ponderosa Pine Ecosystem). Mapped by Kuchler as K- 17 (Ponderosa Pine-Douglas-fir Forest). Interior Douglas-fir forest cover type (SAF 210). No SRM for Douglas-fir cover types in the southwest region. Pseudotsuga mensiesii Association 122.611 in Douglas Fir-White Fir (Mixed Conifer) Series 122.61 of Rocky Mountain Montane Conifer Forest 122.6 of Brown et al. (1998, p. 37). Pseudotsuga mensiesii Series (Ludwig et al., 1986). Riparian vgetation: Willow-Dogwood Series 232.51 in/of Rocky Mountain Swamp and Riparian Scrub 232.5 of Cold Temperate Swamp and Riparian Scrub 232 (Brown et al, 1998, p. 44). Southern Rockies- Sedimentary Mid-Elevation Forest ecoregion 21f (Omernik and Griffith, 2006).

Two types up closer- The range vegetation of a riparian zone along a smaller stream and lower edge or first contact of a second-growth Douglas-fir forest in the Sangre de Cristo Mountains of northern Neew Mexico. The riparian plant community included a stream side shrub zone comprised of red or Pacific willow, thinleaf or mountain alder, water birch, and Rocky Mountain maple and an outer or exterior herbaceous zone dominated by sleepygrass and with cutleaf coneflower as the major range forb species.

These two views are of the same range plant communities introduced in the immediately preceding two slides. Range animals on the grazing land shown here were limited to mule deer (Odocoileus hemionus) and elk (Cervus canadensis), two extremely valuable game species.

While photographing this mesic forest range landscape, the Nikon-toting rangeman was serenaded by black-billed or Amerrican magpies (Pica hudsonia) who were not pleasantly thrilled by human presence intruding into their scenic and otherwise serene home. Unfortunately, these beautiful members of the crow or jay family ( Corvidae) did not shown up distinctly in these wide-angle lens slides.

Simmering at stream's edge- The bank of the Cimmaron River in northern New Mexico provided an example of comparatively species-rich riparian (streamside) range vegetation. Major shrub species were red or Pacific willow, Drummond's willow (Salix drummondiana), thinleaf or mountain alder, water birch, oceanspray or rock-spirea (Holodiscus dumosa), and Rocky Mountain maple.

In addition to providing abundant browse for game species like mule deer and elk, such riparian scrub or shrubland helps provide high-quality habitat for fish and other animal species (vertebrates, including furbearers like beaver (Castor canadensis) and invertebrates) and, perhaps most importantly overall, protection of watershed with assurance of high-quality runoff water for numerous purposes ranging from irrigation to municipal use.

Trees in the forest immedately above the stream in the first slide were Douglas-fir and Rocky Mountain juniper (Juniperus scopulorum).

One species of many- Red or Pacific willow (Salix lasiandra= S. lucida subsp. lasiandra) growing along a stream in the Sangre de Cristo Mountains of northern New Mexico. Red or Pacific willow is one of 28 Salix species (if this author counted correctly) reported for the Land of Enchantent by Allred and Ivey (2012, ps. 531-535). In addition, there were a few subspecies in some of the more widely distributed Salix species.

Willows are some of the most important browse-providing and stream-protecting range plants. The Salix genus is a "taxonomic nightmare" to all but specialists. There are far fewer species of cottonwood (Populus spp.), another genus in family Salicaceae, which are also important (and typically dominant) woody plants of riparian vegetation.

Sleepy by streamside- Sleepygrass (Stipa robusta) shown as several large plants (first slide), basal shoots (second slide) and panicle (third slide) growing in outer (herbaceous) zone of a riparian range plant communityalong a stream in the Sangra de Cristo Mountains of northern New Mexico. The common name of "sleepygrass" is derived from the condition in which horses become lethargic, slow-moving, and dull-acting as if in a stupor as a result of consuming forage of this species. Apparently a toxic substance produced by a fungal endophyte in this grass is responsible for that unresponsive condition (Allred and Ivey, 2012, p. 629). Sleepygrass intoxication was treated in most of the poisonous plant references including Burrows and Tyrl (2013).

The comparatively narrow panicle with ascending branches is characteristic of several Stipa species and was shown to good advantage in the third image.

Sleepygrass was discussed in that ole standby, Range Plant Handbook (Forest Service, 1940, G120). Readers desiring a more recent treatment (though without any management considerations) can find sleepygrass in Barkworth et al. (2007, ps. 129-130) under a newer scientific name (one perhaps based on cladistic criteria which, of course, were rejected by the present author).

Technicl note: the 'bleached" first image was due to malfunction of a Nikoor lens when aged lubricant in the tool prevented proper operation of the shutter. Repair of the lens to fully operationalstatus came too late for this image, but students can tell that the herbaceous zone had developed along the outer edge of the riparian plant community. Fortunately, the lens did function properly for the second and third slide.

Santa Fe National Forest, Taos County, New Mexico. Late July; milk-stage of grain.

Southern and Middle Rocky Mountain Forests - IA
(Including Regionally Related Range Types)

Ponderosa Pine Forests and Associated Types or Subtypes

Rocky mountain-Great Plains Transition

Ponderosa Pine Forests and Adjoining Riparian Vegetation

Southern and Middle Rocky Mountain Forests - IA (Including Regionally Related Range Types)

Ponderosa Pine Forests and Associated Types or Subtypes

Rocky mountain-Great Plains Transition

Ponderosa Pine Forests and Adjoining Riparian Vegetation

Southern and Middle Rocky Mountain Forests - IA
(Including Regionally Related Range Types)