Alpine

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In its descriptions of rangeland cover types the Society for Range Management (Shiflet, 1994) recognized and described only two alpine grazing types outside of the Alaska Region: Alpine Rangeland, SRM 410, in the Great Basin Region and Alpine Grassland, SRM 213, in the Pacific Southwest Region. Beyond doubt, the alpine types of the Rocky Mountains below the Arctic region are distinct and deserving of their own designation and description. In absence thereof, the range vegetation covered in slides below are variants of SRM 213, SRM 410, and, possibly, SRM 902 (Alaskan Alpine Herb). The alpine biome is likewise given but brief coverage in Agriculture Handbook 475 (Garrison et al. 1977) as FRES No. 44, the Alpine Ecosystem. Zwinger and Willard (1972) is an exceptional readily read reference. By far the most relevant publication for consideration of the Rocky Mountain alpine ecosystem is the work reported (Bowman and Seasfedt, 2001) for Niwot Ridge on the Front Range of the Southern Rocky Mountains (Colorado). The description and discussion of the structure and function of the alpine tundra ecosystem edited by Bowman and Seastedt (2001) was especially pertinent for viewers studying examples of alpine range in Rocky Mountain National Park that were presented below. The earlier seminal work by Marr (1961) was recommended as a prelude to the synthesis provided in Bowman and Seastedt (2001).

1. View from a jet-liner of the landscape where the Southern Great Plains (the High Plains= Staked Plains= Llano Estacado) and Black Mesa and Sangre de Cristo Ranges of the Southern Rocky Mountains meet. Range vegetation types go from shortgrass plains and mixed prairie grassland types through the pinjon-juniper woodland type at the foot slopes up through the ponderosa pine forest type to timberline and finally the alpine ecosystem. Union County, New Mexico. February, hiemal aspect.

2. Another aerial view of the Southern Rocky Mountains giving a tansect glimpse of both the Front and Sangre de Cristo Ranges under heavy snow pack. The dark forested areas are the numerous forest range types viewed previously (Douglas fir, ponderosa pine, quaking aspen, Englemann spruce-subalpine fir cover types); the mountain tops, most of which are snow-covered so as to be clearly visible, comprise the alpine range types. Viewed as either ecosystem(s) or landscape these "islands in the sky" are the highest natural pastures, the elevational limit of our native grazing lands. The range types and range sites of the alpine zone, recognized as being unique at least as far back as the life zones of C. Hart Merriam, are habitats differing from other range environments in those abiotic factors associated with high elevation such as more limited oxygen, different qualities and quantities of radiation, extremely diverse and steep relief, etc. Based on such criteria as shortness of growing season, soil features (eg. freeziing/thawing) and microsites of permafrost (permanently frozen soil or soil layers like subsoil or other earthen deposits), and most importantly, similarity or sameness of vegetation the alpine ranges were originally regarded as a kind of tundra. Originally the term alpine as a noun (from the European mountains, the Alps) was not used and instead these "natural pastures in the sky" were treated as a kind of tundra (tundra is Lapp and/or Russian for "marshy, unforested land"). Weaver and Clements (1929, 1938) divided Tundra Climax or The Tundra into Arctic Tundra and Alpine Tundra. The latter in North America was subdivided into Petran Tundra and Sierran Tundra for the alpine climaxes of the Rocky Mountains and Sierra Nevada Mountain. The designation of alpine tundra is still widely used and seems appropriate when describing the vegetation of all those geographic locations that share a common physiogonomy and many of the same plant species, and especially for the vegetation above timberline in the Arctic region. Differences in elevation, limited occurrence of permafrost in the alpine of the Rockys and Sierra, and the greater extent of mature soils in Arctic than in mountain tops (alpine ecosystems) are substantial enough that most contemporary ecologists seem to regard alpine as different from classic tundra (eg. Arctic tundra that has extensive permafrost). Thus these two closely related, perhaps sometimes overlapping, designations are typically viewed as separate biomes or formations, large-scale ecosystems, and range types. The Society for Range Management in its examples of kinds of rangeland has distinguished alpine from tundra since it first defined rangeland in A Glossary of Terms Used in Range Management (Kothmann, 1974).

*Note: In the classification scheme of Brown et al. (1998) Rocky Mountain alpine was shown only at biotic community rank (fourth level) as Rocky Mountain and Great Basin Alpine Tundra with indication that future study could warrent separation of this into two biotic communities. No series (fifth level) distinctions were given. The same obtrained for Cascade-Sierran Alpine Tundra and other North American alpine ecosystems. As such, the Brown et al. (1998) biotic community nomeclature was not shown further below, the specific biotic community being obvious from stated location.

3. The alpine landscape as viewed from the edge of an alpine meadow— The perimeter of this alpine range is dominated by species of dwarf willow (Salix spp. [they are a taxonomic nightmare to key out]). Immediately outward from the willow scrub is the krumholtz (German for "twisted wood") or elfinwood, the highest extension of timberline, the ecotone of woody vegetation where the combination of high winds, bitter cold, and various forms of frozen precipitation prune and deform "stray" individuals from the subalpine forest below. It is also possible that the stunted, deformed trees or shrubs of woody species like subalpine and Englemann spruce are unique ecotypes (genotypes adapted by natural selection to local or distinct habitats) capable of survival at the elevational limits of these forest species. The various elevational forest zones (forest cover types and life zones) can be seen aligned below, beginning with the subalpine fir-Englemann spruce type. Riparian communities of Salix and Populis species outline the aquatic and hydric habitats of the Cache La Poudre River that is the headwaters of the mighty Colorado River which downstream has carved as its own wonderland one of three Grand Canyons in North America. Rocky Mountain National Park, Grand County, Colorado. August, estival aspect. About 12,000 foot elevation.

4. A composite view of Rocky Mountain plant communities going from Alpine Meadow dominated by willow with microsites of grasses and sedges to krumholtz of Englemann spruce through subalpine forest ending with the willow-cottonwood riparian scrub of the Cache La Poudre River. The undulating ground surface is due to solifluction (the flow of water-saturated soil or ground over an impermeable layer of permafrost, bedrock, etc. due to alternate freezing and thawing; in other words, frost-heaving on a larger scale). Solifluction results in various ground surface features like terraces. Frost hummocks also form due to action of frost-heaving. The overall result is a pattern of alternating mounds and depressions over the land. As will be seen below, this patterned ground is very pronounced across the Arctic Tundra. The presence of patterned ground and solifluction terraces in the alpine is evidence of the similarity between alpine and tundra. Rocky Mountain National Park at about 12,000 feet, Grand County, Colorado.

5. Alpine Tundra in the actual or true sense of the term. A layer of permafrost beneath waterlogged soil that alternatively froze and thawed resulted in this solifluction terrace that holds melt water from snow. The terrace and impermeable permafrost pond water and create a micro-habitat that supports willow (the most common alpine species here are Salix brachycarpa, S.planifolia, and S. arctica). In front of the willow is the climax sward of alpine meadow known as Alpine Turf comprised of Carex and Gramineae species. The conspicuous white flowers are of the alpine composite, black-headed daisy (Erigeron melanocephalus). Behind the terrace is a seral stage of alpine vegetation between Alpine Turf and Fellfield (described below). Elevation: 12,500 feet. August, estival aspect. Rocky Mountain National Park, Grand County Colorado. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation for alpine types in the central Rocky Mountains. SRM 410 is the closest approximation. Zwinger and Willard (1972) is an outstanding reference for the Rocky Mountain alpine ecosystem for beginners. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006). Marr (1961) and Bowman and Seastedt (2001) were recommended for advanced students.

6. Timberline Krumholtz or elfinwood of Englemann spruce is the uppermost "outpost" of montane forest of subalpine fir and Englemann spruce from the Hudsonian life zone below. This is a narrow ecotone (a transition zone between adjacent communities, even of biome scale, between the subalpine montane forest and the alpine zone. Pruning, and even killing, by desiccation of spruce is plain). "Flagging" is a telltale characteristic of timberline trees and shrubs. Elevation: 12,500 feet. August. Rocky Mountain National Park, Grand County, Colorado. Timberline vegetation is FRES No. 23 (Fir-Spruce Forest Ecosystem). K-19 (Spruce-Fir-Douglas-fir Forest). SAF 206. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

7. Where wind and relief reign supreme— Timberline and alpine vegetation reflect dominant role of wind (and wind-driven frozen precipitation) along with topography in carving a mosaic of plant life across the landscape of the Colorado Rockys. Note the cirque in the middle ridge line. A cirque is a steep-walled, typically craggy, or cliffed head, of a mountain valley formed by the erosive forces of a glacier; the bowl-shaped depression at the head or, less commonly, along the sides of a glaciated valley. Note also snowbanks which are often actually firn or neve (snow of at least one year of duration that is on its way to becoming glacial ice). Rocky Mountain National Park, Grand County, Colorado. August; estival aspect.

8. Landscape-scale view of the alpine ecosystem- This mountain top shot pictoralized the concept of alpine ecosystems as "islands in the sky". It was, however, a landscape view such that this "top of the Rockys" panorama included both timberline with it's krumholtz form of conifers and the uppermost reaches of the Englemann spruce-subalpine fir forest (non-krumholtz form) as well as the alpine zone. The rounded green spots were the elfinwood (krumholtz) of Englemann spruce and various alpine species of willow. This photograph thus illustrated the concept of Landscape Ecology as the discipline dealing with interconnected ecosystems where the scale of ecosystem was no larger (greater) than that of biomes or biome ecotones (ie. alpine, forest, timberline).

Besides the natural resource professions of Range Management, Forestry, and Wildlife Management this slide illustrated importance of Watershed Management (even in an exteme drought). It "drove home" importance of alpine range as invaluable for watershed.

Rocky Mountain National Park. Elevation of peak was slightly over 12,000 feet. June (but in a hot drought year).

9. Alpine ecosystem-This mountaintop panorama shows the "patchwork " of alpine vegetation in various stages of plant succession due to such things as soil at different degrees of development, micro-relief, animal disturbance, varying exposures to wind, etc. Alpine range is unique in that there is extreme diversity of seral stages within relatively short distances of themselves. Rangemen can stand in one location and study the development of alpine vegetation because the landscape mosaic varies over such short distances that space handily substitutes for time. In addition, at an instant of time an alpine landscape provides a toposequence: an arrangement or pattern of plant communities along topographic gradients like north- vs. south-slopes or higher vs. lower elevation such that degrees of soil development or microsite features either override successional relations among the communities or that represent seral relatedness over extremely long time frames. Alpine vegetation is also a chronosequence: an arrangement of seral plant stages caused by differential states of vegetation recovery from disturbances or even primary plant succession at an instant of time. The typical alpine landscape is a checkerboard-like arrangement of alpine meadow interspersed with fellfield and boulder fields. Fellfield means "rock field" or "stone field" which is alpine land that is 35% to 50% rock-covered and which supports lichens, mosses, and cushion plants (covered below). Boulder fields are similar to fellfield except with larger rocks.

Zwinger and Willard (1972) is an excellent reference for "tenderfeet"; Marr (1961) and Bowman and Seastedt (2001) are of the caliber for mountain rangemen.

Rocky Mountain National Park, Grand County, Colorado. Elevation is 12,500 feet. August, estival aspect. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation for alpine types in the central and southern Rocky Mountains. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

10. Picture-post card scene of the alpine range ecosystem- Tops of the Colorado Rockys provided a backdrop for a fell field in an advanced stage of plant succession. The vegetation featured here was dominated by grasses (especially Poa species) and grasslike plants. Among the latter, Carex species were predominant, especially curly or rock sedge (C. ruprestis), but other genera included Elocharis and, as discussed below, Kobresia. Very few plants of these species were in flower- and fruit-stage and identification by your flat-lander author was not possible. Beidleman et al. (2000) listed 17 Poa species for Rocky Mountain National Park. Some of the most abundant on alpine range included inland bluegrass (P. interior), alpine bluegrass (P. alpina), Greenland bluegrass (P. rupicola), Arctic bluegrass (P. arctica), and mutton grass (P. fendleriana ssp. longiligulata). Beidleman et al. (2000) described over 60 Carex species growing in Rocky Mountain National Park. Such arrays encompassed a diversity of niches and clearly not all would be found in any given local area such as the one shown here.

The nearly total absence of forbs should be noted. Alpine forbs, as explained below, tend to be more abundant (as both number of species and their composition of the vegetation based on cover, density, etc.) on sites and habitats that are less advanced successionally. The successionally and geologically most advanced alpine sites (and vegetation) are the alpine meadows which, as implied by "meadow", are dominated by graminoids (grasses and grasslike plants).

The very tops of peaks in the background are precisely speaking above the alpine. Some of the patches of snow are technically snow beds or even small glaciers (ie. perpetual ice and snow). This the highest elevational zone that is literally above the alpine zone was designated as the nival zone. In the original Life Zone scheme of C. Hart Merriam this elevational (and, on continental-scale, latitudinal) "layer" was the Arctic of the Arctic-Alpine Zone. Even in the nival zone there is vegetation (known as nival flora) but this is generally of the algae group which forms what are known as snowbed communities. These are more common on perennial snow (known as neve). Rangemen and foresters should not consume such water without first boiling it otherwise they can develop "scours".

Rocky Mountain National Park, Larimer County, Colorado. June. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designtions for alpine cover types in the central and southern Rocky Mountains. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

11. An alpine meadow in early to mid-sere. Seral-stage sedges like rock or curly sedge (Carex rupestris) are major soil-forming agents that will improve the sere for more advanced successional species like blackroot sedge (Carex elynoides) and kobresia (Kobresia myosuroides).

Rocky Mountain National Park, Grand County, Colorado. August, estival aspect. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

12. The functioning alpine ecosystem. Elk or wapiti grazing this alpine meadow at what appears to be somewhat mid-sere stage (somewhere between fellfield and the climax sedge-grass turf) and that would seem to be Fair to Good range condition class. Is this simply a stage in development of alpine meadow? Or is it retrogression, range deterioration, due to overgrazing by elk? Or if it is a community induced or maintained by elk that is thus a grazing disclimax? Or is it a zootic climax overriding the climatic climax of Alpine Turf?

Rocky Mountain National Park at 12,000 foot elevation. Larimer County, Colorado. August, estival aspect. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

13. Alpine fellfield. This "rock field" above the trees has progressed to the geologic stage of rock weathering such that the rocks are more the size of gravel than of stones. Some lichens and mosses have persisted from earlier seral stages, but forbs are the dominant category of range plant. Species here include alpine avens (Acomastylis rossii), yellow stonecrop (Sedum lanceolatum), goldflower (Hymenoxys acaulis), alpine clover (Trifolium dasyphyllum), and dwarf clover (T. nanum). This is a successionally advanced fellfield.

Rocky Mountain National Park, Larimer County, Colorado at 12,500 feet elevation. August. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation for alpine types in the central or southern Rockys; parts of SRM 410 are an approximation. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

14. Close-up, detailed view of fellfield— The plant community is less advanced along the path of succession that seen in the preceding view. Here vegetation still consist primarily of cushion plants ( herbaceous, perennial plants that form a dense, convex-shaped, rounded, heavy mass of many leaves and short stems that protrude above the dome-shaped, compact leaf mass thus resembling a pin cushion) and carpet plants (short plants of uniform height spreading by stolons or rhizomes thereby forming carpet-like coverings over the ground surface). Plant species visible include the classic cushion plant, moss campion (Silene acaulis), various clovers (Trifolium spp.), alpine phlox (Phlox condensata), alpine sandwort (Minuartia obtusiloba), alpine nailwort (Paronychia sessiliflora= P. pulvinata), and various stonecrops (Sedum spp.). These plants grow in the boundary layer (the zone between the soil surface and the shallow layer of air that remains relatively calm; the soil is warmer than the calm air just above it). This detail of the plant community illustrates why alpine ecologists often describe the alpine as "deserts atop mountains".

Rocky Mountain National Park, Larimer County, Colorado. August. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation: SRM 410 is closer than SRM 213 to the Rocky Mountain alpine. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

15. Alpine in late spring- Wind-swept ridge of a fellfield composed mostly of cushion plants and carpet plants still in winter dormancy stages even though it was two days before the summer solstice. Dominant (at least the major) species in foreground was moss campion. Other species included those listed immediately above.

Rocky Mountain National Park, Larimer County, Colorado. June. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation: SRM 410 is closer than SRM 213 to the Rocky Mountain alpine. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

16. Scarce groceries for local fauna- Alpine fellfield with rock chuck or yellow-bellied marmot (Marmota flaviventris) foraging for what little feed he can find. Alpine plants were just emerging from (or still in) winter dormancy two days before summer solstice. It had been an unusually cold, wet winter and belated spring from which these alpine plants were emerging, but even in years of milder weather conditions climate at alpine elevations (this sscene was at roughly 11,800 to 12,000 foot elevation) is harsh resulting in extremely short growing seasons. Engelmann spruce-subalpine fir subalpine forests were in the distant background.

Marmots hibernate to get through the long cold alpine winters. Upon emergence from hibernation males start searching for feed and mates. This is a harem-building species with three or four females per adult dominant male. Several of these harems congrete to form small colonies that consist of dens or borrows under boulders and natural rock piles on these mountain top habitats. This male marmot had already been habituated to getting food from photograph-snapping tourists and was not in the least ashamed to come begging (albeit it in a rather low-key, quite dignified manner). When this fellow was disappointed by the free-loading photographer who caught the chuck's winter-leaned conformation the poor begger left disgruntled and disgusted (and apparently muttering something about fat, lazy lowlanders). It was not a "home run" for the photographer either however as he had his 28mm wide angle lens on and the marmot left before a normal lense could be fitted in the bayonet ring of the Nikon FM.

For those sympathsizing with the marmot and suspecting the author of staging the shot with the rock chuck enticed to an area of ususually sparce pickings roll on to the next two slides…

17. Late spring snowbank (it might as well have been the dead of winter)- This was the only alternative to the almost non-existant herbage of still-dormant alpine plants on the fellfield ridges of alpine range introduced above and photographed two days before the summer solstice. At least the hungry marmot presented above could find water. This snowback was in a slight depression on the leeward side of a ridgetop. The range plant community beneath this snowpack was more mesic than that of the drier shallower soils on the ridge. This edaphic-moisture difference resulted in a graminoid (grass and caric sedge)-dominated turf vegetation in contrast to the adjacent graminoid-forb sporadic (interrupted cover) vegetation on the wind-blown and more xeric habitats.

18. Supposed to be summer- Early summer on an alpine range at roughly 12,000 foot elevation. Most of a heavy snowpack had melted, but a few snow banks were still around three weeks into summer. Some Engelmann spruce of the krumholtz form were in the background that marked timberline. Slightly below treeline an ephemeral stream lined with riparian scrub of dwarf willows. Most of the alpine vegetation (that above the krumholtz marked timberline) was either Alpine Meadow or Alpine Turf or a much smaller area of Wet Marsh. These forms of alpine vegetation were treated below where they were presented at other than the landscape-scale used here.

There was also a snow bed community composed primarily of red-pigmented green algae (Chalmydomonas nivalis). This cryophlif algae is responsible for the phenomenon of "pink snow" or "watermelon snow" which, incidentially, is laxative and mildly toxic so high-elevation tramplers are best advised to refrain from eating alpine snow. In the newer taxonomic schemes algae are in kingdom To old school adherents of the "its either in the plant or animal kingdom" (like your crotchy author) C. nivalis "watermelon snow" comprised a distinctive range plant community on the alpine range. In the ecosystem concept C. nivalis is a decomposer or reducer member of the alpine ecosystem.

Rocky Mountain National Park, Larimer County, Colorado. 8 July. For the vascular range plant community of Alpine Turf or Alpine Meadow the following units were the closest fit. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation: SRM 410 is closer than SRM 213 to the Rocky Mountain alpine. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

19. "What were you saying about summer time?"- In the alpine,climate is severe and weather extremely variable. A cobalt-blue sky with nary a cloud in it suddenly "filled to the brim" with an immense thunderstorm. Within a span of a quarter hour (or less) precipitation came as a sequence of cold rain, freezing rain and sleet, hail, grupel and, finally snow (crystalline flakes). Temperature was in range of 41-44 degrees Farenheit. At an elevation of approximately 11,500 feet this alpine fell field appeared as a "winter wonderland" with alpine avens (Geum rossii), old-man-of-the-mountain or alpine sunflower (Rydbergia grandiflora), moss campion (Silene acaulis subsp. subacaulescens) bravely standing like guards on the range top frontline out from under a layer of ice. Alpine avens was far and away the dominant (at least in this late vernal or early estival plant society) while alpine sunflower (the post card flower of the alpine and just about everyone's favorite) was the associate species. Less common alpine range plants included alpine clover (Trifolium dasyphyllum) and pretty or twisted-fruit draba (Draba streptocarpa).

This thunderstorm contained lots of lightning, including the deadly cloud-to-ground lightning. . The photographer "defied death" as much as he dared and then got off the mountain, pronto. Lightning is common in the Rocky Mountains, especially with thunderstorms during the warmer seasons.. Lighning can be particularily dangerous in the alpine because storms form quickly, lightning tends to strike higher objecrts like mountain tops, and mountain visitors are far from shelter. Use extreme caution when in the moutains, especially in the higher alpine and late spring-summer seasons. Human death can come instaneously, literally in a flash.

20. Summer ice in the alpine- A combination of liquid and frozen precipitation (rain, sleet, hail, grupel, and snow) from a fast-developing thunderstorm (complete with flash lightning) and temperatures in the low 40s (degrees Farenheit) left this partial covering of ice on an alpine fell field range (elevation of about 11,500 feet) in early summer. The dominant plant at this later vernal-early estival plant society was alpine avens. Other common range species in their full-bloom stages at time of photographs included alpine sunflower, alpine clover, moss campion, and pretty or twisted-fruit draba.

The latter of these was represented by the middle plant in the second slide. Pretty draba was flanked on the left by alpine avens and moss campion and on the right by moss campion.

Some Notable Range Plants of the Alpine

The species diversity of the alpine range ecosystem is astonding. This is especially the situation for alpine forbs. Not only are there numerous species of forbs on the alpine range but they come from many different families. A sample was selected and presented beginning now. Almost all of these "alpine characters" were growing on a ridge overlooking Forest Canyon in Rocky Mountain National Park at an elevation of about 11,000 feet.

21. Alpine sorrel (Oxyria digyna). A species living in rock habitats ranging from boulder piles to fellfields. Rocky Mountain National Park, Grand County, Colorado at over 12,000 foot elevation. Polygonaceae (buckwheat family). August.

22. Moss campion (Silene acaulis ssp. subacaulescens). This is the archtypical cushion plant in full bloom. Moss campion is a member of the pink family (Caryophyllaceae). A Carex species is "keeping it company" (moss campion is serving as a nurse plant and generally improving the alpine micro-environment for successionally more advanced species). The pavement of gravel and small stones is similar to desert pavement and justifies the description of alpine as "mountain-top deserts" or "desert islands in the sky". Rocky Mountain National Park, Grand County, Colorado. Elevation is over 12,000 feet. August.

23. Parry clover (Trifolium parryi). The classic carpet plant. It too calls fellfields "home". There are several other Trifolium species in the alpine and adjacent range ecosystems. Rocky Mountain National Park, Grand County, Colorado. Elevation is over 12,000. August.

24. Alpine clover (Trifolium dasyphyllum)- There are a number of Trifolium species in the Rocky Mountains. Beidleman et al (2000) described nine species (three alien) of Trifolium in Rocky Mountain National Park. Though small by clover standards this species is one of the most delicately beautiful of all the range forbs. Timberline (about l1,000 foot elevation).

25. Old-man-of-the-mountain (Rydbergia grandiflora= Hymenoxys grandiflora )— One of the most showy and conspicuous plants on the alpine. This "dwarf sunflower" is another widely distributed species growing on habitats ranging from fellfield to climax meadow communities. It is especially plentiful in seral patches transitional from fellfield to the climax Alpine Turf. The common name comes from the long pubescence that diffuses the quantities of ultraviolet light that are twice those at sea level. The genus name is a commenrative one after P.A. Rydberg, Curator of the New York Botanical Garden, who authored some of the earliest and finest florae of Colorado, the Rocky Mountains, and central North American prairies and plains. Overall the Compositae is the number one family of range forbs (on basis of both number of species and biomass), but on alpine range the sunflower family is not nearly as dominant or prominent as on other biomes like grasslands and deserts. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren).

26. Snowball saxifrage (Saxifraga rhomboidea)- The saxifrage family (Saxifragaceae) was represented on the alpine range by this long-stemmed species with the diamond-shaped (ie. rhomboid) basal leaves. It was growing on a microsite still damp from melt water at an elevation of about 11,000 feet. Rocky Mountain National Park. June.

27. Sky pilot (Polemonium viscosum)- Enter now the phlox family (Polemoniaceae). This species is a denizen of fell fields and gravel slopes. It also comes in on disturbed areas like gopher eskers, a phenomenon discussed below. Like most of the other characters in this alpine cast, sky pilot grew on a mountain top at approximately 11,000 feet elevation. Rocky Mountain National Park. June.

28. Alpine phlox (Phlox condensata= P. sibirica= P. pulvinata= P. caespitosa, plus combinations of these at the subspecies level!)- This is the second member of the phlox family (Polemoniaceae) to grace your monitor. This species with it's conspicuous flattened growth form was included as a textbook example of a carpet plant. This delightful specimen was growing at roughly 11,000 feet. It came back out on stage as a component of fell field vegetation below. Watch for it. Rangemen must be observant. Rocky Mountain National Park. June.

29. Alpine or western wallflower (Erysimum capitatum)- This crucifer (member of the mustard family, the Cruciferae or Brassicaceae) grows over a wide range of elevations equally at home in the montane, subalpine and alpine zones.This bright specimen was growing at timberline (approximately 11,000 feet). Rocky Mountain National Park. June.

30. Alpine parsley (Oreoxis alpina)- This alpine range forb respresented the parsley or carrot family (Umbelliferae or Apiaceae) that is distinguished by having the umbel type inflorescence in which all of the pedicels have a common point of origin on the peduncle. Alpine parsley has a spreading, prostrate habit that is a common morpholocigal adaptation of apline plants to the extreme conditions of wind and general xeric habitats of the alpine ecosystem like fell fields. Plants with this general growth form are characterized as carpet plants. Rocky Mountain National Park. June.

31. Alpine forget-me-not (Eritrichum aretioides)- This is another carpet (= mat-forming) plant of the alpine range. It represents the borage family (Boraginaceae). Most borage members have some form of extra-obvious pubescence. This is a survival feature to provide protection from excessive light, transpiration, heat or cold, dessication, etc. in much the same way as spines, hairs, and fluted stems in cactus.

32. Greenleaf chiming bells (Mertensia lanceolata)- This is another member of the borage family. This specimen grew on a fell field at an elevation of about 11,000 feet. Rocky Mountain National Park, Larimer County, Colorado. Early August.

33. Silver scorpionweed (Phacelia hastata)- Presented here is a member of the relatively small Hydrophyllaceae (waterleaf family). Plants of this group have showy-- often coiled or wrapped-- inflorescences with prominent stamen and styles. In some biomes and cover types waterleaf family members grow on fertile and protected sites often serving as field indicators of mesic and moderate habitats. In western North America most of the Phacelia species grow on disturbed, rocky habitats. As indicated by their common names, most are usually thought of as weeds. The individual featured here was growing in an alpine "rockpile" right at timberline (about 11,000 feet). Another Phacelia species was presented as an understorey forb in the limber pine forest (Forests and Woodlands section, Southern and Central Rocky Mountains). Rocky Mountain National Park. June.

34. Alpine avens (Geum rossii= G. turbinatum)- This member of the rose family (Rosaceae, Rosoideae subfamily) is one of the more common and larger forbs in alpine and subalpine zones. Thus, alpine avens comprises a relatively large proportion of the species composition of alpine range on basis of foliar cover and biomass. This species is adapted to various habitats in the alpine ranging from those on gravelly, shallow soils to others having mesic, better-developed soils. According to Beidleman (2000, p. 214) this range forb is a component of the "hay" stored by pika. Rocky Mountain National Park (about 11,500 feet elevation). June.

35. White marshmarigold (Psychrophila leptosepala)- The buttercup family (Ranunculaceae) was represented by this moisture-loving forb that grows in both subalpine and alpine zones. It is probably more common on subalpine mountain meadows especially in depressions where snow melt-water accumulates. That was where this attractive specimen was "caught". White marshmarigold is nonetheless an alpine range forb. It often grows in association with a number of the other species in the "line-up" presented here. Rocky Mountain National Park. Elevation was around 10,000 feet. June.

36. Yellow stonecrop (Sedum lanceolatum)- Now it is time for the stonecrop family (Crassulaceae) to enter center stage. This small, rather obscure family is famous beyond it's size due to discovery of a third photosynthetic pathway in members of this taxon: Crassulacean Acid Metabolism (CAM). Readers are referred to any standard text in Plant Physiology. The references published by the Society for Range Managaement (Sosebee, 1977; Bedunah and Sosebee, 1995) are outstanding. The benefit of CAM, the most water efficient of the three discovered photosynthetic pathways, and it's evolutionary development in alpine as well as desert species attest to the general xeric nature of many habitats on alpine range. Yellow stonecrop grows on shallow, rocky (ie. generally xeric) microsites in contrast to it's mesophytic "cousin" featured next in our alpine cast of characters. Rocky Mountain National Park (11,000 foot elevation). Early August.

37. King's crown (Sedum rosea)- The importance of Crassulacean Acid Metabolism (and the ecophysiological lessons it illustrated) warrented inclusion of another stonecrop. Besides, this one was just "too purty" to omit from our cast of characters. Used in combination with yellow stonecrop this species provided an example of adaptation to habitat and ecological niche. King's crown prefers moist to wet habitats on the alpine range. This contrast with the xeric habitat of the preceding yellow stonecrop. Here was a mountain top illustration of speciation and niche selection. Rocky Mountain National Park at an elevation just below 11,000 feet. Early August.

38. Alpine sandwort (Lidia obtusiloba= Minuartia biflora) - This sterotypic carpet plant is a member of the chickweed family (Alsinaceae) which has often been treated as a subfamily (Alsinoideae) of the pink family (Caryophyllaceae). This is a common species on rock outcrops. It presents a striking appeance, but only when viewed at eye-level. Alpine forbs are wild flowers for native plant "purists" and others not opposed to getting down on hands and knees. It was herein recommended that those who prefer the the easily viewed gaudy cultivars stay out of the hills and leave the native Lilliputian flower gardens to those who appreciate them. Folks wanting merely something that they can see on the fly (and who desire outrageously sized blooms) only crowd the roads and hiking trails for those who truly enjoy the alpine and other native plants. Those who want only big ornamentals that are easily seen while standing up should stay home.

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

39. Alpine bistort (Bistorta vivipara= Polygonum viviparum)- This and the next range forb are in the buckwheat family (Polygonaceae). This first Bistorta species is a high-moisture species preferring depressions or boggy habitats in both subalpine and alpine zones. Flush water conditions or not, wet alpine and subalpine meadows are harsh environments from the standpoint of short growing seasons, cold temperatures, strong dessicating winds, etc. Alpine bistort has so adapted to this habitat that it not only flowers but reproduces asexually by shedding bublets equipped with one new leaf to carry out photosynthesis for the little sister plants falling from the parent plant. Another example of a clonal organism with each bublet-derived plant a ramet of the same genetic individual (the genotype is the genet).

Rocky Mountain National Park, Larimer County, Colorado.Elevation was slightly over 11,000 feet. August.

40. American bistort (B. bistortoides= P. bistortoides)- American bistort is more common than alpine bistort. Both species are found in the alpine and subalpine zones, and both prefer mesic soil conditions though B. bistortoides is not so finicky about it. Both of these species were treated by Dayton (1960, ps. 75-77) as members of the Polygonum genus.

Rocky Mountain National Park. June.

41. Yellow western paintbrush (Castilleja occidentalis)- Even the figwort or snapdragon family (Scrophulariaceae) had members in this alpine cast of characters even though this family is represented by more species on grasslands and deciduous forests. Habitat was a fell field at about 11,000 feet elevation. Rocky Mountain National Park, Larimer County, Colorado. Early August.

42. Esker of northern pocket gopher (Thomomys mazama). Esker is the geologic term for a long narrow sinuous ridge or mound comprised of sand, gravel, and/or boulders deposited by a stream that is flowing in conjunction with (as inside or beneath) a stagnant glacier. This term is employed in a poetic-like usage to describe the winding tunnels made by earth-dwelling rodents like gophers. The impact of this form of herbivory is to induce retrogression in the range plant community (ie. to cause range deterioration) from which vegetation recovers by secondary succession. As with grazing by elk this could be interpreted as either overgrazing or a zootic (=grazing) disclimax. Such disturbances in combination with the patchwork of varying successional stages of vegetation development create on alpine range a landscape mosaic in miniature. Rocky Mountain National Park, Grand County, Colorado. August. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation for alpine types in the central or southern Rocky Mountains (SRM 410 is closest).

43. Fresh (this winter's) gopher esker- Localized disturbance by northern pocket gopher. At the edge of this melting snowbank is the remains of a serpentine or sigmoid-shaped burrow built by the northern pocket gopher. Part of the aboveground tunnel has already collapsed while the part that extended up into a pre-existing snowbank (and still partially supported by it) remained intact. Such rodent activity resulted in considerable soil mixing and was a poignant example of pedoturbation, the general process of mixing in soil. This was a special case of faunal pedoturbation, such mixing by animals (from earthworms and rodents to man, the farmer). Pedoturbation by the native pocket gopher is a natural factor in pedogenesis (soil formation) and, as mentioned in the next caption, a natural disturbance. Such disturbances have taken place for millennia, and vegetation and ecosystems developed and evolved hand-in-glove with them.

Close inspection of this scene revealed that, in spite of soil mixing, impact on existing plants was minimal. Observant viewers will have seen that perennial cespitose graminoids (bunchgrasses, sedges, etc.) were not eaten, dug out, or covered by soil. If proportion of bare ground was increased this merely created seedbeds for establishment of new plants.

Holes dug by gophers contribute to soil aeriation and water infiltration, but without experimental data there was no way to know if this was beneficial or detrimental to plants. For instance, deep burrows (miniture shafts) could in theory (ie. until proven by experiments) direct much precipitation deeper into the soil profile and farther away from the bulk of plant roots. Mere philosophizing or theorizing never proved anything by the standards of science. Only controlled experiments or, at least, simple comparisons can do that. That is the scientific method.

Rocky Mountain National Park (almost 12,000 feet elevation), Larimer County, Colorado. June.

44. Recovery of vegetation on a recent gopher esker- Plants were beginning to invade and revegetate the area of an alpine fell field denuded by northern pocket gophers visible on the left side of this photograph. The portion of land on the right side was not directly impacted by the rodents, but served as a "refuge" or relict area from which propagules (both asexual and sexual) were disseminated to facilitate the processes of secondary plant succession. This was an example of patch dynamics at micro-scale. Such disturbances, especially at this micro-patch scale, maintain species diversity and are but one of many factors of denudation that contribute to the dynamics of vegetation, the phenomena and unifying concept central to development and management of native plant communities.

Rocky Mountain National Park, Larimer County, Colorado. Elevation approximately 12,000 feet. June.

45. Patterned ground of the alpine rangeland. As explained above, the freeze/thaw processes and shrink/swell outcomes taking place in the soil profile cause various arrangements of soil, stones, and other earth materials on the surface of these native grazing grounds. Rock patterns include simple ones like rock streams and rock stripes to ornate geometric shapes like rock garlands and rock polygons. Fragments of soil parent materials and mountain geologic strata such as the granite seen here vary in size from boulders to gravel. A boulderfield complete with boulderpiles is seen here. These arrangements of larger rock are home to the pika (Ochotona spp.), the lagomorph that stores "haystacks" of herbage and various fruits in the rock crevices for winter feed. These arrangements of rock are also home to the hibernating rockchuck or yellow-bellied marmot (Marmota flaviventris).

Rocky Mountain National Park Colorado. August. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation for alpine types in the central or southern Rocky Mountains. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

46. Another view of rock-patterned alpine range- This and the preceding photograph were "cross-sectional" or end-on views of a form of rock patterns like those described immediately above. Vegetation was a mix of various forbs as well as graminoid species. Viewed at microsite scale there were habitats of bare rock, unweathered parent material (granite), surrounded by habitats determined by immature soils at varying stages of pedogenesis (ie. soil development).

The nival zone that was mentioned in the preceding caption was even more evident in the background of this slide (ridgeline on peaks in right background for example).

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

47. Patterned ground form of alpine rangeland near the climax state- This was a lengthwise perspective or a view looking across land having a pattern of rock stripes. It presented a view of rock-patterned alpine range that was roughly a 90 degree difference from that of the two preceding slides.

There were relatively few forbs in the vegetation of this site (in contrast to that of the range community shown in the preceding slide). The most advanced soil and vegetation in the alpine range ecosystem (other than wetland forms) is the Alpine Turf or Alpine Meadow (presented and discussed below). In the classic model of vegetation development (ie. the Clementsian paradigm) Alpine Turf is the climax for alpine. The grass and grasslike plant-dominated range shown here had advanced successionally to stage of subclimax on the theoretical alpine sere that would terminate at climax Alpine Meadow. Interestingly, the soil was not at the state or stage of maturity (the termination of soil genesis) found on Alpine Meadow, but the dominant range species seen here was one of the dominant alpine range plants of the mature stage of soil genesis and the climax stage of vegetation development. The tufted herb with it's telltale mixture of yellowing and still-green shoots was Kobresia myosuroides. This species was presented below.

Rocky Mountain National Park (roughly 11,000 foot elevation), Larimer County, Colorado. June. (This photograph was taken in the summer of the driest one year in Colorado weather records, 2002. Plant phenology had progressed beyond what it is typical for this time in the annual cycle. The climax K. myosuroides had progressed to a phenoloical state typical of late summer.)

48. Landscape and general appearance of alpine range- Shown here was the variation in alpine range vegetation due to local differences in land form and soils on the "islands in the sky". The fell field in the foreground had fairly well-weathered granitic parent material and better-developed soils than thoseof most fell fields. Commensurately, vegetation had developed closer to the climax Alpine Turf community that ultimately forms on mature alpine soils (the Tundra great group of the 1938 soil classification system). The different colors of vegetation indicated various plant species and/or plant communities. For example, light tan or pale yellow zone was vegetation dominated by miscellaneous Carex species. These spots had until recently been covered by snow such as the localized snow banks remaining in the background. Plants in these patches, a high proportion of the cover of which was rock or curly sedge, were recently under snow had barely emerged from dormancy following snow melt and had not yet "greened up". The small patches of dark green color were some species of alpine willow that survived in the microsite of a hillside seep. Evidence of geologic erosion was visible on this steep south slope.

Rocky Mountain National Park, Larimer County, Colorado. Elevation was estended from about 11,000 feet at foreground upward. June. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designations for alpine cover types in the central and southern Rocky Mountains. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

49. Fell field range plant community- Species diversity on alpine range often appears to be greatest on fell field forms rather than on gravelly, Alpine Turf, or Alpine Marsh forms of alpine vegetation. Successional status of fell fields is dependant on the theoretical framework in which it is viewed.

In polyclimax theory fell fields could be viewed as edaphic climaxes, the highest successional development possible on the soil of a given fell field and under the combination of abiotic and biotic factors currently existing. It follows logically that vegetation which developed on even less mature (less developed) soils is also edaphic climax. All such plant communities that are in dynamic equilibrium with average current conditions are the potential natural vegetation (even on bare rock).

By contrast, under monoclimax theory fell fields or any habitat lacking mature soils are a seral stage (more precisely, an array of seral stages) that in climatic age (or perhaps the geologic time of a peneplain) would ultimately develop into the mountain top regional climax of alpine turf (shown below). Mature soils (eg. Alpine Meadow, the intrazonal great group of the 1938 soil classification system; Tundra great group of the zonal soils under the 1938 system) of alpine (= alpine tundra in some descriptions) ecosystems (or landscapes) support meadows of Alpine Turf (= Alpine Meadows).

The immature soils of boulder fields prevents their classification as climax under monoclimax theory, but they are edaphic climaxes under polyclimax theory. Biological diversity has often been shown to be maximum (greatest species richness) at mid-sere or seral stages below that of climax. Such interpretation may be biased somewhat toward monoclimax (or modified monoclimax) theory. It was explained in the accompanying review of climax theories that the polyclimax theory was accepted (by implication if nothing else) in the Range Management profession with adoption of the range site concept. Determination of potential natural vegetation (usually climax vegetation) based on taxonomic units of soils (usually the series or association) was an application of the Tansian polyclimax interpretation. In this practical application of the polyclimax perspective all of the general seral stages in development of alpine vegetation could be interpreted and described as range sites (eg. fell field, mid sere; fell field, complete as well as Alpine Meadow).There could be range site descriptions, including lists of species and their response to disturbances (ie. decreaser, increaser, invader), for seral stages which could be understood as "permanent" in time scale of human management. In fact, this could also be done under rubric of monoclimax theory with the only difference being that Alpine Meadow (= Alpine Turf) would be labeled climax while successional stages still had site descriptions complete with species compositions under various relative degrees of deterioration from the "ideal" or "standard" under management that did not retard plant succession or induce retrogression.

Either way, fell fields are generally the "rock gardens" of the alpine.

This slide presented a headhigh, eye-level view of the Rocky Mountain rock garden of a fell field range as would be seen by a standing rangeman. The white flowers were alpine phlox and the "yeller" ones be alpine parsley. Alpine touch-me-not at pre-bloom stage was common as were Poa, Carex, and Kobresia species (also most at pre-bloom). Granitic parent material provided rock for the range "rock garden". Rocky Mountain National Park at 11,000 feet elevation (approximately). June.

50. Fell field species- Here is a squatting rangeman view of fell field vegetation. It was a portion of the vegetation in the left foreground of the preceding slide. In the field of view presented here alpine phlox was conspicuous along the left hand margin. Alpine clover was also readily discernable in the upper right hand corner by the piece of granite surrounded by and appearing to be engulfed by carpet plants. Yellow flowers of alpine parsley were "keeping company" with the alpine clover. The cespitose grasslike "critter" was one of the most interesting in this cast of alpine characters. More on it immediately below.

Viewers should make special note of the patch of bare soil. The gravelly content of this fell field ground was suggestive of the soil surface of many immature desert soils. Some ecologists, especially ecophysiologists or autecologists, have for instructive and conceptual purposes described the alpine environment as an "edaphic desert". This was in contrast to true (= climatic) deserts. The "edaphic desert" analogy or metaphor was employed to emphasize the fact that immature soils (most fell field soils would be grouped in the Entisol order of the Soil Taxonomy system) in combination with other abiotic factors of the alpine environment result in desert-like conditions for alpine plants. The invention of an instructive term like "edaphic desert" was a useful coinage to impart the concept that much, if not most, of the alpine range ecosystem is generally a xeric, harsh habitat for plants. Over a great part of the alpine range this is a biological reality. It is especially the case on windy ridges kept free of snow, south and west slopes, and stoney sites like boulder fields and scree slopes.

Like the desert, however, there are range sites in the tundra and alpine biomes that have mature soils (eg. Desert and Tundra zonal great groups and Alpine Meadow intrazonal great group of the 1938 soil classification system). Where not impacted by recent or alien forms of disturbance mature soil equals climax vegetation (ie. development of soil and vegetation occur concommitantly and interdependently). The immature soil and seral stage vegetation presented in this slide illustrated that fact, especially when the soil and vegetation components shown here were compared with mature soil and the climax vegetation of Alpine Turf (= Alpine Meadow) and Alpine Marsh presented in slides below.

The sample of alpine range viewed in these last two photographs was at some seral stage of soil and vegetation development below (before or prior to) the terminal stage of general alpine climax and mature soil. This is conceptually true when viewed from the Clementsian (= monoclimax) model. In perspective of the Tanslian polyclimax model the plant community and soil seen here would be viewed as the ecological or natural potential for that specific site (ie. the highest state of development possible now not in the geologic future). In context of the polyclimax model it should be noted that the ecological (= successional) state in this photograph was not in retrogression (not in a state of deterioration) but rather the maximum for a fell field. The question is whether fell field (at highest state of fell field soil and vegetation development) is one kind of climax. Either way, there is a state of soil maturity and expression of vegetation more advanced than that shown here, but the Alpine Meadow and Alpine Marsh (which were covered below) can be viewed as separate range sites (just as could a hypothetical Fell Field, Mid Sere or a Boulder Field be "range sites"). Fell fields would thus be interpreted as another range site (or other range sites if more that one).

Rocky Mountain National Park at about 11,000 foot elevation. June.

51. Turf species in advanced development of vegetation on a fell field- If polyclimax theory was accepted such that there are several range sites or habitats types on alpine range, each with it's potential natural vegetation (ie. many alpine climaxes exist), it could be argued that there is a climax fell field plant community. If there is such a climax fell field vegetation, it would likely closely approximate the sample shown in this photograph which included one of the dominant species of the Alpine Turf. It was explained later in this section that Alpine Turf (= Alpine Meadow) was the mesophytic form of alpine range that developed on mature soil and that has been interpreted as the highest expression of vegetational development in the alpine ecosystem (ie. the climax for the Petran Tundra in the Clementsian monoclimax model).

One of the most diagnostic dominants (an indicator species) of Alpine Meadow is the grasslike plant, bogsedge (Kobresia myosuroids= K. scirpina= K. bellardii= Elyna bellardii). Three plants of this highly tufted species were featured prominently in this slide (the three clumps of combined yellow and green shoots resembling a bunchgrass).This species has been viewed as being one of the dominants of the climax alpine pastures. Weber and Wittmann (2001) described it as "[t]he climax dominant on mature soils of relatively dry but peaty alpine tundra". The common name (most commonly none was given by taxonomists) is somewhat misleading because the most characteristic habitat is not bogs or the alpine wetlands (ie. the Alpine Marsh) of the alpine ecosystem but rather the mesic alpine meadows. Beidleman et al. gave Kobresia myosuroides the common name of "superturf" and declared it "the dominant plant on mature, snow-free areas of tundra that have deep soils". Mutel and Emerick (1992, p. 193) identified "sedge kobresia" as dominant on "level or gently rolling areas with fine-textured, humus-rich, stable, deep soils that are moist through the growing season" adding that Kobresia communities often grow among "ancient patterned ground figures". Kershaw et al. (1998) described it's habitat as "dry, open, wind-blown sites". That was precisely the environment of this advanced fell field plant community though the soil was not deep and rich in humus.

This was yet another example of the importance of microsite (microclimate, ground-level climate, etc.). Quick inspection of this photograph by readers will reveal that the Kobresia plants grew adjacent to granite rocks. These rocks served to shed and direct water and to block drying winds that would increase evaporation of soil water. This combined result due to presence of outcropping rock permitted a mesic (mesophytic) species to grow on a site that was generally more xeric than optimum habitat for the mesophyte. On this fell field weathering of rock and development of soil had progressed to about as great a degree as possible short of the mature soil of the Alpine Meadow form of alpine range.

Rocky Mountain National Park (overlook above Forest Canyon at elevation of roughly 11,000 feet). June.

52. Kobresia myosuroides- These were the two plants in the left midground of the preceding slide. Given the role or ecological status of this tufted grasslike plant as the dominant climax species of alpine meadow range it has remained downright obscure in the literature. In addition, Kobresia species (there are a total of three Kobresia species in the Colorado Rockys) have unique-- and confusing-- floral parts. These members of the Cyperaceae have paired staminate and pistillate florets inside the spikelet (which does hot have the fused perigynium). Until this structural arrangement is understood Kobresia species are easily confused with certain of the Carex species. Plus, there is considerable variation in the habit (perhaps phenotypic plasticity) of K. myosuroides (which stands to reason given the commonness and locally wide distribution of a dominant plant). On the windy overlook where these specimens grew their culms were considerably flattened such that even at over eight inches in length plant height above ground was less than half of that. K. myosuroides is a tufted plant, but it is not strictly cespitose as it has short rhizomes. The rhizomatous feature in combination with prolific tillering often creates the mat-like habit shown here. Interestingly, the most consipuous, readily observed feature of K. myosuroides is the combination of green culms and leaves along with yellow ones and/or yellowed bases of growing shoots.

Rocky Mountain National Park (elevation of 11,000 feet on an overlook of Forest Canyon), Larimer County, Colorado. June.

53. Alpine turf developing on a fell field- Turf dominated by Kobresia myosuroides forming on a highly weathered fell field. Various species of sedges and grasses (especially bluegrass species) were also plentiful. Forbs were present but they were uncommon. The forb component lacked both the dominance and species diversity of less successionally advanced fell field vegetation shown above (ie. "farther down" the sere from the Alpine Turf climax). The vegettion presented here should be compared that on fell fields having less mature (less-developed) soils and then to the climax vegetation of mature soils shown below. This array from bare rock to climax Alpine Meadow (Tundra great group soils) provided the student with the successional sequence of the general sere of the "alpine tundra" or what ecologists like Weaver and Clements (1938, p. 481, 485-486) described as Petran Tundra (Carex-Poa association). In context of species diversity (= biological diversity, often the clipped "biodiversity") the following statement was summative of the alpine range vegetation displayed throughout this section: "… alpine societies are best developed in subclimax of disturbed areas, in which the control of the dominant sedges and grasses is not yet complete. In the ultimate consociation of Elyna bellardi [the current Kobresia myosuroids], for example, the dominance of this sedge is so complete as practically to exclude all but a few individuals of the forbs that thrive in the more open communities of Carex rupestris and the grasses" (Weaver and Clements, 1938, ps. 485-486). The alpine community presented here was at roughly the subclimax stage and had a diverse mixture of forbs ranging from alpine clover to alpine avens as well as rock sedge and Kobresia myosuroides.

Rocky Mountain National Park, Larimer County, Colorado. Elevation: about 11,000 feet. June. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine meadows and Barren). No SRM designations for alpine cover types in the central and southern Rocky Mountains. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

54. Pyrenanean sedge (Carex pyrenacia), known also as Crandall sedge (C. crandallii)- The climax alpine plant community of Alpine Meadow is dominated by grasses and grasslike plants. The two major families of grasslike plants in the alpine ecosystem are the same two as on most other biomes: sedge (Cyperaceae) and rush (Juncaceae). One of the more common and productive sedges is the species presented here. It has a solitary, linear spike inflorescence and and broad, robust leaves which often persist into the next growing season. It is a cespitose species with lower leaves that are fairly well developed. This specimen was growing along the moist edge of a solifluction terrace. Elevation slightly below 12,000 feet. Rocky Mountain National Park, Larimer County, Colorado. Early August.

55. Alpine transition sere community. This seral stage is between fellfield and the alpine climax meadow. This vegetation is dominated by curly sedge and American bistort (Bistorta bistortoides), but a large portion of plant cover is still cushion and carpet plant species. The immature soil is between the lithosols of fellfield and the Alpine Meadow great group (the 1938 zonal-intrazonal-azonal classification system). FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designation specific for the Rocky Mountain alpine but some of the description in SRM 410 might help. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

56. The alpine climax. Alpine Meadow or Alpine Turf and its accompanying mature soil, Alpine Meadow (the intrazonal great group of the 1938 Soil Classification System). Grasses and sedges common to Alpine Turf include numerous bluegrasses such as Arctic (Poa arctica), alpine (P. alpina), and Greenland bluegrass (P. glauca); tufted hairgrass (Deschampsia caespitosa); numerous sedges such as blackroot sedge, cliff or Rocky Mountain sedge (Carex scopulorum), hair sedge (C. capillaris), and man-hater or shortleaved sedge (C. misandra); and bog-sedge (Kobresia sibirica). In classic succession theory this is the mesophytic (=climatic= regional) climax of the Rocky Mountain alpine range type.

Rocky Mountain National Park, Grand County, Colorado. August at an elevation over 12,000 feet. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barren). No SRM designtion for alpine types in central or southern Rockys, but SRM 313 for alpine in the northern Rocky Mountains and SRM 410 for Basin Range, Wasatch, and Unita Series described similar alpine communities. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

57. Alpine Marsh. The hydrophytic form of climax alpine complete with a mature soil that by 1938 Soil Classification System is classified as the Tundra great group. There is a permafrost layer in this soil that is responsible for the ponded water which is held back by a solifluction terrace formed by the freezing-and-thawing and alternating shrinking and swelling of this mature zonal soil. This plant community is almost exclusively of Carex species especially Rocky Mountain sedge, Nelson's sedge (C. nelsonii), and water sedge (C. aquatilis). The scattered plants bearing conspicuous white inflorescences atop bare stalks are American bistort.

Rocky Mountain National Park, Colorado at elevation of about 12,000 feet. August. FRES No. 44 (Alpine Ecosystem). K-45 (Alpine Meadows and Barrens). No SRM designation specific for this site or phase of alpine, but some of the description/discusssion in SRM 410 seemed similar. Closest fit in (Brown, 1998, p. 39) was Mixed Bunchgrass Series 141.21 of Rocky Mountain Alpine and Subalpine Grassland biotic community 141.2. Southern Rockies- Alpine Zone Ecoregion, 21a (Chapman et al., 2006).

58. Alpine-timberline ecotone of the Northern Rocky Mountains— Intricate pattern of the krumholtz form of subalpine fir-whitebark pine (Pinus albicaulis)-Englemann spruce "interwoven" with climax Alpine Turf of blackroot sedge, alpine bluegrass, and glacier lily (Erythronium grandiflorum). Logan Pass (elevation, 6600 feet), Glacier National Park, Montana. July. Alpine typically occurs at lower elevations in the northern than in central or southern sections of the Rockys, undoubtedly a latitude X elevation interaction. Northern Rocky Mountain alpine— Alpine Meadow dominated by glacier lily (left) and climax alpine turf of blackroot sedge and alpine bluegrass. The bare or more open space in center foreground is an area disturbed by Columbian ground squirrel (Spermophilus columbianus) causing alpine turf to retrogress back to the pre-climax alpine meadow stage. Subalpine fir and whitebark pine are attempting an uppermost invasion here at elevation just above timberline. Logan Pass (6600 foot elevation), Glacier National Park, Montana. July. FRES No. 44 (Alpine Ecosystem). Canadian Rockies- Crestal Alpine-Subalpine Zone Ecoregion, 41b (Woods et al., 2002).

59. Northern Rocky Mountain alpine- Alpine meadow dominated by glacier lily (left) and climax alpine turf of blackroot sedge and alpine bluegrass. The bare or more open space in center foreground is an area disturbed by Columbian ground squirrel (Spermophilus columbianus) causing alpine turf to retrogress back to the pre-climax alpine meadow stage. Subalpine fir and whitebark pine are attempting an uppermost "toe-hold" here at elevation just above timberline. Logan Pass (6600 foot elevation), Glacier National Park, Montana. July. FRES No. 44 (Alpine Ecosystem).Canadian Rockies- Crestal Alpine-Subalpine Zone Ecoregion, 41b (Woods et al., 2002).

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