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USDA United States Department of Agriculture Vegetation of the
Forest Service Rocky Mountain Glacier Lakes Ecosystem Research Station Fort Collins, Colorado 80526 Experiments Site Research Paper RMRS-RP-1 Claudia M. Regan Robert C. Musselman June D. Haines Abstract
Regan, Claudia M., Robert C. Musselman, and June D. Haines. 1997. Vegetation of the Glacier Lakes Ecosystem Experiments Site. Research Paper. RMRS-RP-1. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky.Mountain Research Station. 36 p.
Vegetation at the Glacier Lakes Ecosystem Experiment Site, a 600 ha research site at 3200 to 3500 m elevation in the Snowy Range of southeastern Wyoming, was categorized and described from an intensive sampling of species abundances. A total of 304 vascular plant taxa were identified through collection and herbarium documentation. Plots with tree species were separated from those without tree species for ordination and classification analyses. Detrended correspondence analysis was used to order plots along major axes of composition variation, which are inferred moisture and topographic gradients. Cluster analysis was used to categorize plots based on composition similarity. The resulting groups were named according to species dominants. We identified and described in detail 4 meadow, 4 thicket or scrub, 3 krummholz, and 2 forest plant associations. Key words: alpine vegetation, subalpine vegetation, plant associations, cluster analysis, floristics, Wyoming, Snowy Range, Medicine Bow Mountains
The Authors
Claudia M. Regan was an ecologist at the Rocky Mountain ~esearchStation. Robert C. Musselman is a research plant physiologist at the Rocky Mountain Research Station. June D. Haines is a biological technician at the Rocky Mountain Research Station.
Publisher
Rocky Mountain Research Station Fort Collins, Colorado February 1998
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Claudia M. Regan. Robert C . Musselman. and June D. Haines
Contents
Introduction ......
Study Area ......
Physiography and Geology ......
Climate ......
Landscape Characteristics ......
Wildlife and Domestic Grazers ......
Current Management ......
Methods ......
Field Methods ......
Floristics Documentation ......
Analytical Methods ......
Results and Discussion ......
Vegetation ......
Plant Associations ......
Conclusions ......
Literature Cited ......
Appendix 1 ......
Appendix 2 ......
Vegetation of the Glacier Lakes Ecosystem Experiments Site
tosociological survey of the spruce-fir forests of the Medi- cine Bow Mountains. Their study provides general in- Introduction formation on the composition and structure of these for- ests. However, their study sites did not encompass the full The Glacier Lakes Ecosystem Experiments Site range of environmental variability in subalpine forests, (GLEES) (figure 1)was established in the Snowy Range of and their results do not indicate a comprehensive floristics the Medicine Bow Mountains of Wyoming to study the survey because only 46 species of vascular plants were effects of air pollution on the structure and function of identified. Billings and Bliss (1959) studied snowbank subalpine and alpine ecosystems (Musselman 1994). Re- environments and vegetation. They concluded that snow- search at the site uses an integra ted ecosystem approach to melt in alpine areas greatly influences floristics and pro- assess the environmental impact of human-induced atmo- ductivity. Several other studies investigated the relation- spheric deposition and climate change in a landscape ship between productivity of the Medicine Bow alpine typical of high elevation wilderness systems. These assess- and the environment (Scott 1963;Smithand Johnson 1965; ments require baseline data to quantify the current envi- Smith 1969). Several species-level physiological studies ronmental and biological status of the ecosystem and to have been done on alpine plants of the area (Mooney and study each biological and physical component. Our study Billings 1960,1965; Mooney 1961; Scott et al. 1970). documents the vascular flora of the entire GLEES and The original vegetation classification (Wirsing 1973; categorizes and describes the vegetation types based on Wirsing and Alexander 1975)focused on forest vegetation quantitative data. This quantitative information is useful and identified 5 habitat types of the Medicine Bow Moun- as a baseline for intensive studies of ecosystem processes tains. Alexander et al. (1986) strengthened the classifica- at the site. tion by again focusing on forested habitat types of this Several researchers have studied the vegetation of the area. Although this study identifies 16 habitat types, it Snowy Range. Oosting and Reed (1952) reported a phy- does not encompass all of the environmental variation
Figure 1. Geographic location of the Glacier Lakes Ecosystem Experiments Site.
USDA Forest Serv~ceRes. Pap. RMRS-RP-1. 1998 present in the forests of the GLEES and does not include The floristics survey documents the occurrence of vascu- information on nonforested areas. lar plant species of the GLEES. The vegetation classifica- A preliminary assessment of the vegetation of part of tion identifies vegetation types that are ecologically dis- the GLEES was conducted using a descriptive, qualitative tinct and occupy a significant area. The plant community approach (Simmons 1994).Landscape habitats of the Lost classification provides a framework to study the biotic Lake, West Glacier Lake, and East Glacier Lake portions of components of the GLEES systems. In addition, the classi- the study area were identified using limited ground recon- fication provides supplementary information to resource naissance, air photo interpretation, and a physiognomic mangers about subalpine and alpine meadow and shrub- classification of the vegetation types. Twelve vegetation/ dominated vegetation. landscape types were identified, mapped, and described based on interpretation of 1:6000 aerial photographs (Simmons 1994). Although the description does not in- clude a comprehensive floristic documentation or a char- acterization of plant communities, it does provide a refer- Study Area ence point and a baseline for the intensive study of the GLEES vegetation presented here. The objectives of this study were to: The GLEES is a 600 ha alpine and subalpine research study area in the Snowy Range of the Medicine Bow 1. Provide a comprehensive, quantitative character- Mountains of southeastern Wyoming. The area is part of ization of the GLEES flora; the Medicine Bow National Forest, which is 55 km west of 2. Identify and describe the composition and struc- Laramie, Wyoming, and 15 km northwest of Centennial, ture of the plant associations of the GLEES; Wyoming, at 41" 22'30" latitude and 106" 15'30" longitude (figures 1 and 2). Portions of the GLEES are mapped on 4 3. Determine the similarity between the GLEES veg- U.S. Geological Survey (USGS) 7.5 minute series topo- etation and the prevalent alpine and subalpine graphic quadrangles: the Medicine Bow Peak, Sand Lake, vegetation of the central Rocky Mountains; and Morgan, and Centennial quads. The high elevation of the 4. Provide baseline information for identifying fu- GLEES is defined by the watersheds of 3 glacial cirque ture ecosystem change. lakes: Lost Lake, West Glacier Lake, and East Glacier Lake.
SAFETY SHELTER o 1 mi N + 1 I I 0 NADP SITE Figure 2. The Glacier 1 km 0 METEOROLOGICAL TOWER Lakes Ecosystem 0 SNOWY RANGE OBSERVATORY Experiments Site re- PRECIPITATION COLLECTOR search study area in = PARSHALL FLUME A NDDN SITE southeast Wyoming (from MEADOW Musselman 1994). The elevation of the GLEES ranges from 31 80 m near Little Brooklyn Lake to 3494 m at Glacier Peak on the Snowy Range ridge.
2 USDA Forest Service Res. Pap. RMRS-RP-1. 1998 The low elevation of the GLEES is east of Brooklyn Lake 1963 ). Much of the study area is at the subalpine to alpine and Little Brooklyn Lake, within the Nash Fork watershed transition. Many perennial and intermittent streams drain of the Little Laramie River. The GLEES elevation ranges the watersheds, and the lower elevation portion of the from 3180 m near Little Brooklyn Lake to 3494 m at Glacier study area is scattered with several perennial and ephem- Peak on the Snowy Range ridge. eral ponds. The forest/alpine, forest/subalpine, montane grassland, and terrestrial/aquatic ecotones dominate the GLEES landscape, which makes it an ideal location to Physiography and Geology study ecosystem dynamics resulting from climate change.
The Snowy Range is underlain by Precambrian gneiss and igneous rock (Thornbury 1965).The range rises from Wildlife and Domestic Grazers the western flank of the Laramie Basin and reaches its highest point on Medicine Bow Peak (3,658 m); the North The GLEES, within the Libby Flats grazing allotment Platte Valley is the western border of the range. The (USDA Forest Service 1985),has been grazed primarily by northern part of the range is characterized by Pleistocene sheep since the late 1800s.Recently the area was restricted glacial drift (Houston 1968) that results in numerous from further sheep grazing. streams and lakes; the GLEES is within this glaciated Several wildlife species also have an influence on the portion. A predominant geologic feature of the GLEES is GLEES vegetation. The dominant ungulate in the area is a ridge of Medicine Peak quartzite intruded by mafic dikes the mule deer, and an occasional elk is sited. Pocket that compose 15 to 20 percent of the bedrock in the Glacier gophers create significant soil disturbance that influences Lakes and Lost Lake drainage basins (Rochette 1994). plant community composition and structure. There have Typical of much of theGLEES area above the lakes, soils been no studies of the fauna at the GLEES. formed from the quartzite bedrock are extremely stony (Hopper and Walthall 1994) and are shallow and mini- mally developed in areas of rock outcrops, talus slopes, Current Management and below the permanent snowfield. High elevation soils are primarily Cryochrepts with some Cryorthents, and The GLEES is part of the USDA Forest Service's Laramie low elevation soils are Cryochrepts and Cryumbrepts. Ranger District of the Medicine Bow National Forest. Cryoboralfs are associated with areas of glacial till. Recreation, the primary use of the area, includes fishing, hiking, overnight camping, motorized snow machines, and skiing. Recreational impacts include secondary trails Climate and occasional campfire rings. Although closed to new mining claims, active claims exist. Most mining claims are Minimum and maximum hourly average temperatures unworked and the only evidence of past mining activity is range from -23 "C to -1 OC in winter, and -7 "C to 21°C in an occasional old exploratory soil pit. There are no signs of summer. Most precipitation is snow that covers most of recent mining activity at the GLEES. The study area is the study area from late October until July. Precipitation roadless but is bordered on the southwest by a Forest ranges from 90 to 125 cm; mean maximum snow accumu- Service road; accessibility is provided by a Forest Service lation is about 2 m. A predominant climatic feature of the trail and several unimproved trails. Off-road vehicular study area is the high wind speed that averages 34 km/ hr; travel is restricted to motorized snow machines. winds are primarily northerly and westerly (Musselman 1994).
Landscape Characteristics Methods
The landscape is characterized by steep, highly dis- sected, rocky terrain. The complex terrain influences local Field Methods wind patterns and snow distribution. Differing wind ex- posure and variable snow depth and duration create mi- A stratified, random approach was used to locat e the crohabitats of varying growing season duration and avail- sample sites. A 100 m2 grid was drawn over a map of the able moisture levels, which are the primary limiting fac- study area. Within each grid cell, another grid of points at tors for plant establishment and growthat high elevations 10 m intervals was drawn resulting in 100 possible sample (Billings and Bliss 1959; Johnson and Billings 1962;'Bliss points per cell. One point from each 100 m2 grid was
USDA Forest Service Res. Pap. RMRS-RP-1. 1998 randomly selected and mapped as the sampling location and Moss (1983). Voucher specimens of all vascular plant for that grid cell. The sampling locations were physically species documented were deposited at the Rocky Moun- located using the grid maps, aerial photography (1:24000), tain Herbarium. and an altimeter. Plot locations were marked on the aerial photographs. Markers labeled with the plot number were established as the center for circular macroplots. Plot Analytical Methods centers were sometimes adjusted 1 to 3 m within the area to be sampled to ensure homogeneous environment and Floristics Summary and Biogeographic A nalysis vegetation based on field observations. This adjustment avoided sample plot overlap of 2 distinct habitats. Less A species presence list was constructed after identifica- than 5 percent of the plots required such adjustment. tions were confirmed. Biogeographic affinities were deter- A 0.01 ha (11.28 m diameter) circular macroplot was mined by consulting standard regional floras (Haines et al. used to measure tree and advanced regeneration strata, 1994).Several major biogeographic groups are represented presence of all vascular plant species, and physical envi- in the GLEES flora; Alpine, Arctic-Alpine, Boreal-Mon- ronment information. Trees (defined as woody plants tane, Great Plains, Montane, and ubiquitous. Secondary 2 1.4 m tall) were measured for diameter at breast height biogeographic groups were also identified; those of par- and the species was recorded. Trees < 1.4m were tallied by ticular interest included Circumpolar, North America, species. All vascular plant species in the macroplot were Western North America, Rocky Mountain, and Southern listed. Physical environment information included eleva- Rocky Mountain. These groups are similar to those sum- tion, aspect, slope position, and slope form. marized by Thorne (1972) and discussed in Brown and A modified Daubenmire (1959) technique was used to Gibson (1983).We have added resolution in biogeographic measure the canopy cover of all vascular plant species. group identification to address all biogeographic distribu- Transects originating from the plot center and extending tions cited in the regional floras. The percentage of flora to the perimeter of the plot were randomly located using falling into each biogeographic group was calculated. azimuth corrected for declination (13"). Subplots (.lm2) were systematically placed at 1 m intervals along the Summary of Vegetation Data transect. A canopy cover class for each species was deter- Detrended correspondence analysis (DCA)(Gauch 1982; mined for each subplot. The cover classes and associated Peet et al. 1988) was used to determine patterns in the percent cover quantities are: 0 ( USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Sokal 1973) and the resemblance function used was per- duce a classificationsystem applicable to the entire GLEES cent similarity (Bray and Curtis 1957). Cluster analysis study area. results were summarized using dendrograms. The spe- cific groups or communities were identified subjectively to achieve definable, interpretable, and repeatable asso- ciations based on our field observations and knowledge of the species characteristics (Ludwig and Reynolds 1988). Results and Discussion The classification analyses were done on the individual data sets identified from the ordination results. Clustering was done by sites and species. Diversity measures were Vascular plant taxa present at the GLEES are listed in calculated and used to compare community structure Appendix 1. A total of 289 species and 304 taxa were (Ludwig and Reynolds 1988). Hill's indices (1973) were documented through collection and herbarium verifica- used to determine the total number of species in the tion. Of the 304 taxa at the GLEES, 102 were not docu- sample (NO),the effective number of species including the mented in the sample stands. Eleven taxa are present in number of abundant species (Nl),and the number of very >40 percent of the macroplots. They are, in order of fre- abundant species (N2). The modified Hill's ratio (Hill quency, Abies lasiocarpa uar. lasiocarpa, Erigeron peregrinus, 1973; Alatalo 1981) was used as a measure of evenness. Hieraciurn gracile uar. gracile, Picea engelrnannii, Vacciniurn Plant community types or plant associations were named scoparium, Potentilla diuersifolia var. diuersifolia, Carex rossii, according to either dominant and codominant species for Polygonurn bistortoides, Sibbaldia procumbens, Senecio nonforested types or dominant overstoryspecies and domi- dirnorphophyllus uar. dirnorphoyhyllus,and Achillea millefoliurn nant shrub or herb layer species for forested types (those uar. lanulosa (Appendix 2). Of the 304 taxa, 91 are present where tree species are present). Where possible, the termi- in < 5 percent of the total number of macroplots. Geo- nology for types followed that used in other published graphic distributions for species at the GLEES are summa- reports. The potential for the use of species as indicators rized in table 1and figure 3. Data on geographic distribu- was analyzed by graphing the centroid distributions of tions are missing for 7 of the 304 taxa because we were individual species on the ordinations. Then, constancy unable to find reference to their distributions in the litera- and frequency data for each species in each type were ture. The GLEES species have predominantly Montane evaluated to identify species with the highest constancy distributions with primarily western North America af- values for a dominance type and the lowest frequency finities. Alpine and Boreal-Montane geographic groups values between dominance types. The classification re- are approximately equally represented in the GLEES flora. sults were field tested, refined, and synthesized to pro- Again, western North America affinities predominate. Figure 3. Geographic distributions of the vascular plant taxa of the Glacier Lakes Ecosystem Experiments Site. Secondary Geographic Group B Other Southern Rocky Mountain Rocky Mountain Westem North American North American Circumpolar Alpine Arctic- Boreal- Montane Great Ubiquitous Alpine Montane Plains Primary Geographic Group USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Only a minor portion of the GLEES flora (~5%of the total Table 1. Summary of the geographic distributions of the flora) is represented by species having a Rocky Mountain vascular plant taxa of the Glacier Lakes Ecosystem Experi- or southern Rocky Mountain range restriction. ments Site. The DCA ordination of all sites (figure 4) indicated the need to separate forested stands from all other stands for Number % of better resolution of the forest gradient. Since this analysis of total is based on floristic composition where the response vari- Geographic distribution taxa flora able is species cover, forested stands are those with tree species present. Stand structure is not considered in this Major group Secondary group initial data division; the forested data set includes areas Alpine Circumpolar 6 2 dominated by erect trees and areas dominated by trees in North American 2 c1 the krummholz form. All subsequent analyses were per- Western North American 34 11 formed on the forested and nonforested groups sepa- Rocky Mountain 6 2 rately. The DCA ordination of meadow and shrub-domi- Southern Rocky Mountain 7 2 nated stands is in figure 5a; species are ordinated in figure Other 4 1 5b and the centroid locations of several key species, either ~~~~i~-Al~i~~~i~~~~~~l~~ 2 1 7 those that are dominant or those that frequently occur, are North American 5 2 indicated by arrows. Similarly, forested stands and spe- Western North American 8 3 cies are ordinated in figures 6a and 6b, and key species Rocky Mountain 0 0 centroids are indicated by arrows in both figures. Southern Rocky Mountain 0 0 Ordinations of meadow and shrub-dominated plots Other 8 3 have axis 1and 2 eigenvalues of .92 and .65 (figures 5a and Boreal-Montane Circumpolar 12 4 5b), and ordinations of forest plots have axis 1 and 2 North American 12 4 eigenvalues of .72 and .57 (figures 6a and 6b). The high Western North American 22 7 eigenvalues for the ordinations of nonforested plots (fig- Rocky Mountain 0 0 ure 5a) indicate that these gradients account for more of Southern Rocky Mountain 0 0 Other 7 2 the variation in floristic comvositionI than in the forested plots. In addition, the length of the axis 1 gradient in Montane Circumpolar 20 7 figures 5a and 5b for the nonforested plots is over 2 times North American 26 9 that of the forested plots (figure 6a and 6b). Finally, the Western North American 70 24 pattern of sample distribution indica tes that the nonforested Rocky Mountain 3 1 plots (figure 5a and 5b) are related more clearly to 2 axes Southern Rocky Mountain 0 0 of variation than are the forested plots (figure 6a and 6b). Other 14 5 In this paper, the ordinations area backdrop for an overlay Great Plains Circumpolar 1 <1 of the classification results and illustrate the pattern of North American 0 0 vegetation distribution in ordination space. Western North American 1 <1 Rocky Mountain 0 0 Southern Rocky Mountain 0 0 Other 0 0 7 Ubiquitous or 8 3 - meadow or shrub-dominated plots disturbed V1 * 1 krurnmholz or forest plots 2 6- 0 c m m0 :E0 rn 'Tg 5- m j m 8,; 4- C w 0 0 m 0 w c 0a E.E 3- 3 c a " :Figure 4. Detrended correspondence analysis ordination of kg !#*3 S .a all sample locations, Glacier Lakes Ecosystem Experiments 0 : 2- w > Site. Axes 1 and 2 are the major axes of variation in sample 7-J a II f3 composition. Eigenvalues are .87 for axis 1 and .62 for axis X*x 2. The response variable is species mean cover so that the ) , , , , , , , , I ordination is based on floristic composition and does not , 6 7 8 lo incorporate forest stand structure. Forested and krummholz plots are those where tree species are present. All other Detrended Correspondence Analysis Scores \ Axis 1 Variation in Species Composition plots are either meadow or shrub-dominated associations. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Figure 5a. Detrended correspondence analysis ordination of all meadow and shrub- dominated plots, Glacier Lakes Ecosystem Experiments Site. Axes 1 and 2 are major axes of species composition variation in the nonforested samples; axis 1 is an inferred Plant Association moisture gradient, with samples occupying drier sites on the left and wetter sites on the SAPUCAAQ right. Eigenvalues are .92 for axis 1 and .65 D SAPL-SABRICALE for axis 2. Plant associations are indicated. a DECEICANI SAPUCAAQ is the Salix planifolia/Carex V DAlNlPODl aquatilis thicket. SAPL-SABRICALE is the A VASCIJUDR Salix planifolia-S. brachycarpa/Caltha leptosepala thicket. DECEICANI is the + GEROICARO Deschampsia cespitosa/Carex nigricans X SACA-SABRICASC meadow. DAINIPODI is the Danthonia 0 CAPUICARU intermediaPoten tilla diversifolia meadow. VASCIJ U DR is the Vaccinium scoparium/ Detrended Correspondence Analysis Scores Juncus drummondii dwarf scrub. GE ROI Axis 1 Variation in Species composition CARO is the Geum rossii/Carex rossii meadow. SACA-SABRICASC is the Salix cascadensis-S. brachycarpa/Carex scopulorum thicket. CAPUICARU is the Calamagrostis purpurascens/Carex rupestris meadow. Figure 5b. Detrended correspondence analysis ordination of species in meadows and scrub or thicket vegetation, Glacier Lakes Ecosystem Experiments Site. Centroid locations of selected species, either those that are dominant or are frequently occurring, are .. - indicated by arrows. Species are: CAAQ is t Carex aquatilis, CA LE is Caltha leptosepala, CAAQ CAN1 is Carex nigricans, CAPU is Calamagrostis purpurascens, CARO is Carex rossii, CARU is Carex rupestris, CASC is Carex scopulorum, DA l N is Danthonia intermedia, DEC E is Deschampsia cespitosa, GERO is Geum rossii, JUDR is Juncus drummondii, PO Dl is Potentilla diversifolia, SABR is Salix brachycarpa, SACA is Salix cascadensis, SAPL is Salix planifolia, and Detrended Correspondence Analysis Scores VASC is Vaccinium scoparium. Axes 1 and 2 Axis 1 Variation in Species Composition are major axes of species composition variation in the nonforested samples; axis 1 is an inferred moisture gradient, with the driest end of the gradient on the left. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Figure 6a. Detrended correspondence analysis ordination of all forest and krummholz plots, Glacier Lakes Ecosystem Experiments Site. Axes 1 and 2 are major axes of species composition variation in the forested samples; axis 1 is an inferred moisture gradient, with drier sites on the left and wetter sites on the right. Eigenvalues are .72 for axis 1 and 57 for axis 2. Plant associations are indicated. Plant Association ABLA-PIENICALE is the Abies lasiocarpa- Picea engelmanniiKaltha leptosepala forest. V ABLA-PIENICALE ABLA-PIENIJUCO is the Abies lasiocarpa- 1 A ABLA-PIENNUCO Picea engelmannii/Juniperus communis I + ABLAIVASC krummholz. ABLAIVASC is the Abies + U A 0 I X ABLA-PIEN/CARO lasiocarpa~Vacciniumscoparium forest. ABLA- 0 o ABLA-PIENISAPL PI ENICARO is the Abies lasiocarpa-Picea --2! 2 4 6 engelmannii/Carex rossii krummholz. ABLA- Detrended Correspondence Analysis Scores PI EN/SAPL is the Abies /asiocarpa-Picea Axis IVariation in Species composition enge/man#ii/Sa/ixplanifolia krummholz. Figure 6b. Detrended correspondence 0 0 0 analysis ordination of species occurring in 0 forests or krummholz, Glacier Lakes Ecosys- DAlN tem Experiments Site. Centroid locations of dominant or frequently occurring species are indicated by arrows. Species are: ABLA is 0 B Abies lasiocarpa, ARC0 is Arnica cordifolia, 0 PONE 0-VASC A LI ARUV is Arctostaphylos uva-ursi, CALE is 0-JUCO '='on 0 o >PIEN Caltha leptosepala, CAN l is Carex nigricans, CARU is Carex rupestris, DAl N is Danthonia intermedia, DECE is Deschampsia cespitosa, H IGR is Hieracium gracile, JUCO is Juniperus communis, JU DR is Juncus drummondii, PI EN is Picea engelmannii, PI FL is Pinus flexilis, PONE is Poa nervosa, RIM0 is Ribes montigenum, SAPL is Salix planifolia, VACE is Vaccinium cespitosum, and VASC is Vaccinium scoparium. Axis 1 and 2 are major Detrended Correspondence Analysis Scores Axis 1 Variation in Species Composition axes of species composition variation in forested samples; axis 1 is an inferred mois- ture gradient, with the driest end of the gradient on the left. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 The cluster analysis of the meadow and shrub-domi- is A bies lasiocarpa-Picea engel mann ii/Caltha leptosepala. Gen- nated plots is summarized in figure 7. Eight plant associa- eral characteristics and associated species are in table 5; tions are recognized based on the degree of clustering, mean percent cover or basal area for trees and diversity reflective of the degree of floristic similarity, field observa- estimates are in tables 6 and 7. tions, and tabular comparisons. The associations are ar- ranged in ordination space along inferred moisture and Vegetation elevation gradients (figure 5a). High elevation associa- tions are: Calamagrostis purpurascens/Carex rupcstris, Geunz Vegetation of the GLEES and its spatial pattern on the rossiilCarex rossii, and Salix cascadensis-Salix brachycarpal landscape is typical of the subalpine/alpine ecotone in the Carex scopulorum. Mid to low elevation xeric to mesic central and southern Rocky Mountains. Coniferous for- associations are: Vacciniunz scopariunz/Juncus drummondii ests dominate the landscape at the low portions of the and Danthonia intermedia1Potentilla diz~ersifolia.Mid to low study area and krummholz stands of these forests domi- elevation wet associations are: Deschanzpsia cespitosalcarex nate at high elevations or exposed sites. The forests often nigricans, Salixplanifolia-Salix brachycarpalCa1tha leptosepala, occur as islands or ribbons and are interspersed with and Salix planifolialCarex aquatilis. General characteristics nonforested sites dominated by meadow or shrub-domi- and associated species are in table 2; mean percent cover nated communities. Drainages and topographic depres- and diversity estimates are in tables 3 and 4. sions have by willow shrub communities or graminoid The forest and krummholz classificationis illustrated in dominated meadows. Areas of heavy snow accumulation figure 8. Most of the forest plots are classified as the Abies and prolonged snow cover have willow shrub types or lasiocarpa/Vacciniunz scoparium type. High elevation mesic grass-sedge meadows. Alpine tundra vegetation occurs to xeric associations are: Abies lasiocarpa-Picea engelmanniil above treeline onexposed, windswept slopes. Much of the Salix planifolia, Abies lasiocarpa-Picea engel nzannii/Juniperus landscape is scree slopes and rock outcrops that are mini- conznzunis, and Abies lasiocarpa-Picea engelnzanniilCarex rossi. mally vegetated, and water or permanent snowfields that The mid to low elevation riparian or wetland association are without vegetation. Figure 7. Dendrogram of meadow and shrub-dominated CAPUICARU communities, Glacier Lakes Ecosystem Experiments Site. The dendrogram illustrates clusters of similar samples and is SACA-SABW hierarchically arranged to reflect different degrees of similar- CASC ity. Clustering is by weighted averages with unweighted pair- grouping, and the measure of resemblance is percent CEROICARO similarity. Plant associations identified with each cluster are indicated. SAPUCAAQ is the Salix planifolia/Carex aquatilis thicket. SAPL-SABRICALE is the Salix planifolia-S. VASCJJ UDR brachycarpa/Caltha leptosepala thicket. DECEICAN I is the Deschampsia cespitosdCarex nigricans meadow. DAINI DAlNlPODl POD l is the Danthonia intermedia/Potentilla diversifolia meadow. VASCIJUDR is the Vaccinium scoparium/Juncus drummondii dwarf scrub. GEROICARO is the Geum rossii/ Carex rossii meadow. SACA-SABRICASC is the Salix cascadensis-S. brachycarpa/Carex scopulorum thicket. DECElCAlYl CAP UICA RU is the Calamagrostis purpurascens/Carex rupestris meadow. SAPL-SABW CALE SAPLICAAQ USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Table 2. General characteristics of meadows and shrub dominated associations, Glacier Lakes Ecosystem Experiments Site. Plant association Associated species Habitat characteristics Calamagrostis purpurascens/ Erigeron pinnatisectus alpine sites, exposed rocky ridge tops and Carex rupestris Festuca brachyphylla saddles, no persistent snowcover, shallow soils CAPUICARU Geum rossii Haplopappus pygmae us Minuartia rubella Paronychia pulvinata Phlox pulvinata Selaginella densa Trifolium dasyphyllum Salix cascadensis- Festuca brachyphylla alpine sites, protected rocky ledges and Salix brachycarpd Geum rossi depressions, prolonged snow cover Carex scopulorum Selaginella densa SACA-SABRICASC Geum rossii/ Artemesia scopulorum subalpine sites to near treeline, prolonged snow Carex rossii Erigeron melanocephalus cover, well-drained soils GEROICARO Erigeron peregrinus Erigeron simplex Lewisia pygmaea Poa cusickii Poa reflexa Polygonum bistortoides Potentilla diversifolia Sibbaldia procumbens Vaccinium scoparium/ Carex rossii subalpine upland sites, prolonged snow cover, Juncus drummondii Erigeron melanocephalus very well-drained soils VASCIJUDR Erigeron peregrinus Erigeron simplex Erythronium grandiflorum Penstemon whippleanus Poa cusickii Senecio crassulus Sibbaldia procumbens Danthonia intermedid Agoseris glauca subalpine upland mesic sites, often in forest Potentilla diversifolia Arenaria congesta openings, prolonged snow cover DAINIPODI Artemesia scopulorum Carex microptera Erigeron peregrinus Minuartia obtusiloba Phleum alpinum Poa reflexa Sibbaldia procumbens Trifolium parry; Deschampsia cespitosal Arnica mollis subalpine wet sites, alluvial benches or protected Carex nigricans Caltha leptosepala areas with prolonged snow cover DECEICANI Carex aquatilis Carex scopulorum Erigeron peregrinus Festuca brachyphylla Juncus drumondii Kalmia microphylla Phleum alpinum Salix planifolia Vaccinium cespitosum Salix planifolia- Calamagrostis canadensis subalpine wet sites, drainages, depressions, and Salix brachycarpd Carex aquatilis seeps, poorly drained soils Caltha leptosepala Carex atrata SAPL-SABRICALE Saxifraga odontoloma Sedum rhodanthum Senecio triangularis - Trollius laxus Continued on next page 10 USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Table 2. ( Cont'd.) Plant association Associated species Habitat characteristics Salix plani folial Caltha leptosepala subalpine very wet sites, stream bottoms or Carex aquatilis Carex canescens depressions with the water table at or near SAPUCAAQ Eleocharis pauciflora the surface Saxifraga odontoloma Sedum rhodanthum Figure 8. Dendrogram of forest and krummholz communities, PLOTW I Glacier Lakes Ecosystem Experiments Site. The dendrogram ABLA-PIEN/ PLOT035 I UP. PLOT072 [ PLOTOY I illustrates clusters of similar samples and is hierarchically arranged to reflect different degrees of similarity. Clustering is by weighted averages with unweighted pair-grouping, and ABLA-PIEN/ CAR0 I the measure of resemblance is percent similarity. Plant PLO lQ15 1 r PLOT124 J PLOT0IO PLOW - PIENKALE is the Abies lasiocarpa-Picea engelmanniYCaltha PI.OTII8 leptosepala forest. ABLA-PI EN/JUCO is the Abies PLOm - PLOTllS -- lasiocarpa-Picea engelmannii/Juniperus communis krumm- PLUI013 PLO ID2 l holz. ABLAJVASC is the Abies lasiocarpa/Vaccinium PLOT056 scoparium forest. ABLA-PI ENICARO is the Abies lasiocarpa- PI.OT~I~ PLOT109 J - Picea engelmannii/Carex rossii krumm holz. AB LA-PI EN/ PI.OT027 PLOlMl SAPL is the Abies lasiocarpa-Picea engelmannii/Salix PLOT13V planifolia krummholz. PI.OT029 - PL01136 - PI.OTII0 PLOT127 I, - I PI.OT117 PLOTl3S - PLOT043 PLOT137 - p1.0 ro36 PLOT122 I PLOT154 PLOT055 PLOT1211 PL01076 ABLANASC PI OTI4I PLOT I03 PLOT155 PI.OTQ17 PLoT0la PI.OTIM PLOT054 Pl.OTn95 1 - PI.01120 PLOIUb7 PLOTOY PL.OTl3I PLOT137 PLor1a PLOTlOl Pl.OTll6 1,- I PLOT142 PI.OTI3S PI.OT144 PLOT149 ABLA-PIEN/ CALE r USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Table 3. Mean percent cover of major species by dominance types derived from COMPAH (Combinatorial, Polythetic, Agglomerative, Hierarchical clustering) (Boesch 1977), classification of meadows and shrub dominated associations of the Glacier Lakes Ecosystem Experiments Site. pp pp- - - Dominance types SACA- SAPL- CAPUI SABRI GEROI VASCI DAlNl DECEI SABR SAPLl CARU CASC CAR0 JUDR POD1 CAN1 CALE CAAQ Calamagrostis canadensis Calamagrostis purpurasce Caltha leptosepala Carex aquatilis Carex canescens Carex nigricans Carex rossii Carex rupestris Carex scopulorum Danthonia intermedia Deschampsia cespitosa Eleocharis pauciflora Erigeron peregrinus Erigeron simplex Festuca brachyp hylla Geum rossii Juncus drummondii Phlox pulvinata Potentilla diversifolia Salix brachycarpa Salix cascadensis Salix planifolia Senecio triangularis Sibbaldia procumbens Trifolium dasyphyllum Trollius laxus Vaccinium cespitosum Vaccinium scoparium N3 n species was absent * mean cover < 0.5% number of stands number of subplots CAPUICARU = Calamagrostis purpurascens/Carex rupestris SACA-SABRICASC = Salix cascadensis-Salix brachycarpa/Carex scopulorum GEROICA RO = Geum rossii/Carex rossii VASCIJ UD R = Vacciniurn scopariurn/Juncus druhmondii DAINIPOD I = Danthonia intermediaPotentilla diversifolia DEC EICAN I = Deschampsia cespitosaKarex nigricans SAPL-SABRICALE = Salix planifolia-Salix brach~carpaKalthaleptosepala SAPLICAAQ = Salix planifolia/Carex aquatilis USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Table 4. Diversity estimates for each meadow and shrub dominated type of the Glacier Lakes Ecosystem Experiments Site. Dominance type NOf N12 ~2~ E4 CAPUICARU SACA-SABRICASC GEROICARO VASCIJUDR DAINIPODI DECEICANI SAPL-SAB RICALE SAPLICAAQ -- -- I average number of species occurring in the sample plots average number of abundant species average number of very abundant species evenness CAP UICA RU = Calamagrostis purpurascens/Carex rupestris; SACA-S AB RICASC = Salix cascadensis-Salix brachycarpa/Carex scopulorum; GE ROICARO = Geum rossii/Carex rossii; VASCIJ UD R = Vaccinium scoparium/Juncus drummondii; DAINIPODl = Danthonia intermedia/Potentilla diversifolia; DECEICANI = Deschampsia cespitosaGarex nigricans; SAPL-SABRICALE = Salix planifolia-Salix brachycarpaXaltha leptosepala; SAP L/CA AQ = Salix planifolia/Carex aqua tilis Table 5. General characteristics of forests and krummholz associations of the Glacier Lakes Ecosystem Experiments Site. Plant association Associated species Habitat characteristics A bies lasiocarpa- Picea Carex rupestris krummholz forest in sheltered engelmannii/Salix plani folia Carex scopulorum microsites on exposed windswept ABLA-PIENISAPL Erigeron peregrinus slopes, prolonged snow cover Kalmia microphylla Salix brachycarpa Vaccinium cespitosum A bies lasiocarpa- Picea Calamagrostis purpurascens krummholz forest on dry, gentle engelmannii/Juniperus communis Carex rossii slopes ABLA-PIENIJUCO Ribes montigenum Salix brachycarpa Vaccinium cespitosum A bies lasiocarpa-Picea Carex rossii krummholz forest on dry, exposed engelmannii/Carex rossii Hieracium gracile steep slopes, sparse understory ABLA-PI ENICARO Juncus drummondii Sibbaldia procumbens Vaccinium scoparium A bies lasiocarpa- Picea Carex illota streambanks, terraces, and engelmannii/Caltha leptosepala Carex microptera topographic depressions, moist ABLA-PIENICALE Carex nigricans to wet sites Deschampsia cespitosa Saxifraga odontoloma Trollius laxus A bies Iasiocarpa~Vaccinium A rnica cordifolia widespread in a variety of upland scoparium Carex rossii sites, mesic to xeric with long ABLAIVASC Erigeron peregrinus snow duration Picea engelmannii Poa nervosa Ribes montigenum USDA Forest Service Res. Pap. RMRS-RP-1 . 1998 13 Table 6. Mean percent cover, or basal area (m2/ha) for trees, of major species by dominance types from COMPAH (Combinato- rial, Polythetic, Agglomerative, Hierarchical clustering) (Boesch 1977), classification of forest and krummholz types of the Glacier Lakes Ecosystem Experiments Site. Dominance types ABLA- A B LA- ABLA- ABLA- PlENl PIEN1 PlENl PlENl ABLAI SAPL CAR0 JUCO CALE VASC A bies lasiocarpa Picea engelmannii Total mean basal area Arnica cordifolia Calamagrostis purpurasce Caltha leptosepala Carex nigricans Carex rossii Carex rupestris Deschampsia cespitosa Erigeron peregrinus Geum rossii Hieracium gracile Juncus drummondii Juniperus communis Kalmia microphylla Poa cusickii Poa nervosa Poa re flexa Ribes montigenum Salix brachycarpa Salix planifolia Trifolium dasyp hyllum Trifolium parryi Trisetum spicatum Trollius laxus Vaccinium cespitosum Vaccinium scoparium N3 n4 mean cover < 0.5% species was absent number of stands number of subplots AB LA-P IE NISAP L = A bies lasiocarpa-Picea engelmannii/Salix planifolia ABLA-P IENICARO = A bies lasiocarpa-Picea engelmannii/Carex rossii A BLA- PI ENIJ U CO = A bies lasiocarpa- Picea engelmannii/Juniperus communis AB LA-PI ENICALE = Abies lasiocarpa-Picea engelmannii/Caltha leptosepala ABLAJVASC = Abies lasiocarpa/Vaccinium scoparium USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Plant Associations Table 7. Diversity estimates for each forest and krummholz type of the Glacier Lakes Ecosystem Experiments Site. Calamagrostis purpurascens/ Carex rupestris Meadows Dominance type NO1 N1 ~2~ E4 The Calamagrostis purpurascens/Carex rupestris (CAPU/ ABLA-PIENISAPL 14.0 7.2 5 .O 0.6 CARU) plant association (figure 9) occurs at the highest elevations and on the most exposed sites at the GLEES. ABLA-PIENICARO 1 1.4 7.7 6.1 0.8 This is an alpine type that grows above timberline on rocky ABLA-PI ENIJUCO 8.7 5.2 4.1 0.7 ridge tops and saddles in shallow soils without persistent snow cover. Environmentally adjacent associations are ABLA-PIENICALE 20.0 11.5 8.4 0.7 Salix cascadensis-Salix brachycarpa/Carex scopzrlorunz, which A BLAIVASC 6.4 3.6 2.9 0.7 occurs on protected sites with more snow cover, and Geunz rossiilCarex rossii, which occurs at low elevations. This type average number of species occurring in the sample plots is characterized by grasses, sedges, and forbs with tufted average number of abundant species or cushion growth forms. Associated graminoids include average number of very abundant species Festuca brachyphylla, and associated forbs include: Erigeron evenness pinnatisectus, Geum rossii, Haplopappus pygmaeus, Minuartia rubella, Paronychia pulvinata, Phlox pulvinata, and Trifoliunz ABLA-P IENISAPL = Abies lasiocarpa-Picea engelmannii/ Salix planifolia dasyphyllum. Selaginella densa is very important in terms of total cover. The CAPU/CARU association is included in ABLA-PI ENICARO = Abies lasiocarpa-Picea engelmannii/ the cushion plants/stone pavement landscape habitat Carex rossii described by Simmons (1994) for the GLEES. ABLA-PI ENIJUCO = Abies lasiocarpa-Picea engelmannii/ The Carex rupestris/Trifolium dasyphyllum (CARU/ Juniperus communis TRDA) plant association in north central Colorado (Hess ABLA-PI ENICALE = Abies lasiocarpa-Picea engelmannii/ 1981; Wasser and Hess 1982) is similar in species compo- Caltha leptosepala sition and topographic location to the above type. Species A B LAIVASC = Abies lasiocarpa~Vacciniumscoparium in the CARU/TRDA type that do not occur at the GLEES Figure 9. The Calamagrostis purpurascens/Carex rupestris (CAPU/ CARU) plant associa- tion occurs at the highest elevations anc on the most exposed sites at the Glacier Lakes Ecosystem Experiments Sites. USDA Forest Service Res. Pap. RMRS-RP-1 . 1998 15 are Arenaria fendleri (Eremogonefenlderi), Lidia biflora, and tion is included in the rock outcrop landscape habitat Oreoxis alpina. Other floristicallysimilar plant associations described by Simmons (1994). Although this landscape reported in the literature include Carex rupestrisllidia biflora type represents over 8 percent of the area surveyed by (Komarkova 1976,1988;Baker 1983) in north central Colo- Simmons, the area vegetated by the SACA-SABR/CASC rado, west central Colorado,and north central New Mexico; type is much smaller since the type is limited to sheltered Carex rupest rislKobresia nzyosuroides (Komarkova 1988)in microsites. west central Colorado; and Calamagrostis purpurascens/ The Salixphylicifolia spp.planifolia/Carexscopulorun~ (Salix Helictotrichon nzortonianum (Komarkova 1988) in west pIanifolia/Carex scopulorunz) plant association is a wide- central Colorado. Several species occur at the Colorado spread type throughout central and western Colorado and New Mexico sites that do not occur at the GLEES (Komarkova 1976, 1988; Webber et al. 1976; Hess 1981; including: Helictotrichon mortonianunz, Carex elynoides, Wasser and Hess 1982). This type, similar to the GLEES Castilleja occidentalis, Trifolium nanum, Potentilla uniflora, type floristically and in terms of elevation and topographic Rydbergia grandiflora, and Kobresia myosuroides (in addi- position where it occurs, is richer in species composition. tion to those listed previously). In addition, the occurrence of Salix cascadensis and the absence of Salix planifolia in the GLEES type are conspicu- Salix cascadensis-Salix brachycarpa/ ous differences. Carex scopulorum Thickets The Salix cascadensis-Sal ix brachycarpalcarex scopulorurn Geum rossii/Carex rossii Meadows (SACA-SABR/CASC) association is in the alpine environ- The Geum rossiilcarex rossii (GERO/CARO) meadows ment on topographically protected depressions and on or (figure 10) occur over most of the alpine to subalpine near rocky ledges. Compared to the CAPU/CARU domi- transition environments at the GLEES. These habitats nated sites, the SACA-SABR/CASC habitats are moister experience moderate snow accumulation and a slightly and have a shorter growing season due to snow accumu- prolonged snow cover, but the soils are well-drained and lation and prolonged cover. The community is also not as the wind-exposed sites dry quickly. This type is character- diverse asCAPU/CARU, havingonly Festuca brachyphylla, ized by grasses, sedges, and forbs without shrub or tree Geunz rossii, and Selaginella densa as important associates species. Associated graminoids include Poa cusickii and with the willow shrub and sedge dominants. This associa- Poa reflexa; and associated forbs are: Artemesia scopulorum, Figure 10. The Geurn rossii/Carex rossii (GE ROICA RO) meadows occur o\~er most of the alpine t 0 subalpine transitio nal environments at tt-le Glacier Lakes Ecosys- tern Experiments Sites. 16 USDA Forest Service Res. Pap. RMRS-RP-1 . 1998 Erigerorz nzelanocephalus, Erigeron peregrinus, Erigeron sinz- (ACRO/TRDA) plant association is documented on the plex, Lewisia pygnzaea, Polygonunz bistortoides, Potentilla Medicine Bow National Forest (Smith 1969).This associa- diversifolia, and Sibbaldia procunzbens. Simmons (1994) de- tion shares some floristic similarity to the minor compo- scribes the GERO/CARO association in the diverse nents of the GLEES type but the dominant species are meadow landscape habitat. different. In addition, the ACRO/TRDA type is found on Several Geum rossii dominated plant associations are wind exposed sites with little snow accumulation, while documented in Colorado and Wyoming. The Aconzastylis the GLEES type is found on sites with prolonged snow rossii/Bistorta bistortoides (Geunz rossii/Polygonunz bistortoides) cover. type is widespread throughout central and western Colo- rado (Sheperd 1975; Spencer 1975; Komarkova 1976,1988; Vaccinium scoparium/Juncus drummondii Hess 1981; Hess and Wasser 1982). Lloydia serotina and Dwarf Scrub Castilleja occidentalis occur in this type and are absent from the GLEES. The Aconzastylis rossiilCarex rupestris plant The Vaccinium scopariurn/Juncus drumnzondii (VASC/ association occurs in north central and west central Colo- JUDR) association (figure 11) occurs across all elevations rado (Hess and Wasser 1982). Trifoliunz nanunz and Lidia in the subalpine environments at the GLEES. This type is biflora occur here but are absent from GLEES. At GLEES, on upland sites with a late-melting snow cover where the Carex rossii has replaced Carex rupestris as the codominant soils are very rocky and well-drained, and the sites dry graminoid. The Aconzastylis rossiil Trifoliunz dasyphyllum quickly. This association is floristically similar to the GERO/ CAR0 type except for the shrub component dominated by Vaccinium scoparium, which does not occur in the meadow community. The VASC / JUDR association is also floristically similar to the Abies lasiocarpa/Vaccinium scoparium forest type. However, except for an occasional individual, trees are absent from the VASC/ JUDR habitats. Also, most descriptions of ABLA/VASC lack Juncus drummondii, which is typically found in meadows. Associ- a ted graminoids of the VASC / JUDR associa- tion include Carex rossii and Poa cusickii; associ- ated forbs are: Erigeron melanocephalus, Erigeron peregrinus, Erigeron simplex, Erythronium grandiflorum, p ens tern on whippleanus, Senecio crassulus, and Sibbaldia procumbens. This type is included in the meadow landscape habitat de- scribed by Simmons (1994). The Vaccinium scoparium-Vaccinium cespitosurn/lidia biflora plant association of north central and west central Colorado (Komarkova 1976,1988) is very similar to the VASC/JUDR type at the GLEES. Both types occur on areas of prolonged snow cover. Lidia biflora does not occur at the GLEES and Juncus drummondii is a codominant with Vacciniumscoparium at GLEES. Figure 1 1 . The Vaccinium scoparium/Juncus drummondii (VASCIJUDR) association occurs across all elevations in the subalpine environ- ments at the Glacier Lakes Ecosystem Experi- ments Sites. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Danthonia intermedia/Potentilla divers;folia environments at the GLEES on alluvial benches, protected Meadow areas, and gently sloping terrain with heavy snow accu- mulation and prolonged snow cover. Soils are flooded by The Danthonia in ternzedialPotentilla diuersifolia (DAIN/ the snow accumulation and are poorly-drained. This type PODI) meadows are found throughout the subalpine and is characterized by a dominance of graminoids including: low alpine environments among krummholz islands at Carexaquatilis, Carexscopulorurn, Festuca bmchyphylla,Juncus the GLEES. The association is generally on gentle, pro- drumondii, and Phleum alpinum. Important forbs include tected terrain with prolonged snow cover; it is often found Arnica nzollis, Calfha leptosepala, and Erigeron peregrinus. in forest openings. These sites are more mesic than those Shrubs are of minor importance and are Kalmia nzicrophylla, occupied by the GERO/CARO and VASC/ JUDR commu- Salix planifolia, and Vaccinium cespitosum. These wet mead- nities. Graminoid associates are Carex nzicroptera and Poa ows are included in the wet meadow/willow shrub land- reflexa.Important forbs in this community include: Agoseris scape habitat described by Simmons (1994). glauca, Arenaria congests, Artemesia scopulorunz, Erigeron The Deschampsia cespitosalCaltha leptosepala (DECE/ peregrinus, Minuartia obtusiloba, Phleunz alpinunz, Sibbaldia CALE) plant association is found throughout the Front procumbens, and Trifoliunz parryi. Again, Simmons (1994) Range and in west central Colorado. Except for the ab- included this in the meadow landscape habitat. This asso- sence of Carex nigricans and the occurrence of Senecio ciation is documented in north central and west central crocatus, Erigeron eximius, and Carex festiuella, DECE/CALE Colorado (Komarkova 1976; Hess and Wasser 1982). The is floristically and topographically similar to the GLEES other locations are very similar floristically and topo- type. A widespread Descahampsia cespitosalCarex spp. type graphically to the GLEES sites. A major difference is the has been documented on the Medicine Bow National occurrence of Solidago spa thulata at the Colorado sites, Forest (Knight and Thilenius 1975). None of the descrip- which does not occur at the GLEES. tions document the codominance by Carex nigricans. How- ever, Carex nigricans is a dominant associated with Juncus Deschampsia cespitosaCarex nigricans Meadow drurnmondii in a Carex nigricans/Juncus drummondii asso- Deschampsia cespitosalcarex nigricans (DECE/CANI) wet ciation, which is a widespread type that is somewhat meadows (figure 12) are found throughout the subalpine similar to the DECE/CANI type at the GLEES (Johnson Figure 12. Deschampsia cespitosdcarex nigricans (DEC El CANI) wet meado\INS are found through1out the subalpine enviiron- ments at the Glacie r Lakes Ecosystem Experiments Sites 18 USDA Forest Service Res. Pap. RMRS-RP-1. 1998 and Billings 1962; Spencer 1975; Komarkova 1976, 1988; Salix planifolia/Carex aqua tilis Thicket Rottman and Hartman 1985). Juncus drunzrnondii is more important in the CANI/JUDR association than in the The Salix planifo1ialCarex aqua tilis (SAPL-CAAQ)asso- GLEES type. ciation (figure 13) occurs throughout the subalpine range at the GLEES in depressions or stream bottoms. These are Salix plani folia -Salix brachycarpa/ the wettest terrestrial sites at the GLEES, with very poorly Caltha leptosepala Thicket drained soils and a water table at or near the surface most or all of the growing season. The shrub component of the The Salix planifolia-Salix brachycarpalCaltha leptosepala community occurs patchily on slightly better drained (SAPL-SABR/CALE) association occurs throughout the microsites. Forbs associated with these shrub patches in- subalpine range at the GLEES on wet sites in seeps, clude Caltha leptosepala, Saxifraga odontolonza, and Sedunz depressions, drainages, and valley bottoms. The soils are rhodanthum. Interspersed among the shrub patches are poorly drained and are moist to wet throughout the grow- microsites with standing water that are vegetated with ing season. The type is shrub dominated and occurs as a only a few graminoid species. In addition to Carexaqua tilis, mosaic of shrub patches interspersed with graminoid Carex canescens and Eleocharis pauciflora are found on these dominated meadows such as the DECE/CANI commu- very wet microsites. nity. At a finer scale, the shrub patches occur on raised The Salix planifolia/Carexaquatilisplant associa tion (may hummocks and alternate with poorly drained forb domi- be Salix phylicifolia spp. planifolia/Carex aqua tilis in the nated patches. Important graminoid associates include literature) is widespread in Colorado and Wyoming Calamagrostis canadensis, Carexaquatilis, and Carex a trata. Associated forbs are: Saxifraga odontoloma, Sedunz rhodanthum, Senecio trinagularis, and Trollius laxus. Simmons (1994) describes a wet meadow/willow shrub landscape habitat, which is comparable to the shrub domina ted microsites, and a tall forb wet meadow, whichis comparable to the forb dominated microsites. The Salix phylicifolia spp. planifolialcaltha leptosepala (Salixplanifolia/Calthaleptosepala) plant association in west central Colorado (Wasser and Hess 1982; Komarkova 1988) is similar ex- cept for the occurrence of Pseudocymopterus montanus and Carex festivella, which are not present at the GLEES. The Salix phylicifolia spp. planifalia/Deschampsia cespitosa (Salix planifolial Deschanzpsia cespitosa) plant association of north- ern Wyoming (Johnson and Billings 1962) and north central Colorado (Hess 1981; Wasser and Hess 1982) is also similar to the GLEES. How- ever, Deschampsia cespitosa is a codominant in the SAPL/DECE type and is much less abundant in the willow dominated wetlands at the GLEES. Other floristic differences include the occurrence of Cas tilleja septentrionalis and Senecio crocatus in the SAPL/DECE type. Figure 13. The Salix planifolia/Carex aquatilis (SAPL-CAAQ) association occurs throughout the subalpine range at the Glacier Lakes Ecosystem Experiments Sites. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 (Hanna 1934; Bierly 1982; Hess 1981; Wasser and Hess shrub with Salix planifolia. Other important shrubs are 1982; Komarkova 1988; Cooper 1990). Befula glandulosa Kalnzia nzicrophylla and Vaccinium cespitosunr. Associated occurs in this type in several other locations but was not species include Carex rupestris, Carex scopulorum, and recorded at the GLEES. Erigeron peregrinus. Simmons (1994) includes all krumm- holz associations as krummholz conifer landscape habitat. A bies lasiocarpa-Picea engelmannii/Salix planifolia The Abies lasiocarpa-Picea engelnzannii/Salix planifolia as- Krummholz sociation is a common timberline type. This type is similar The Abies lasiocarpa-Picea engelnzannii/Salix planifol in to vegetation identified as Abies lasiocarya-Picea (ABLA-PIEN/SAPL) krummholz association (figure 14) engelnzannii/Salix y lauca, Picea engelnzannii/Salix is common in the alpine range at the GLEES. These forests pseudolapponunz, or Picea engelnzannii-Abies lasiocarpa/Sal ix occur in sheltered microsites with prolonged snow cover pseudolapponurn in the literature (Sheperd 1975; Hess on exposed windswept slopes. The stands occur as patches 1981; Hess and Wasser 1982; Komarkova 1988). Species surrounded by more exposed areas that are dominated by that are common or characteristic of the type in other areas CAPU/CARU or GERO/CARO associations. Tree species but are absent at the GLEES include Vacciniunz rnyrtillus, and shrubs are both dominant and reflect a matted, Betula glandulosa, and Polemonium pulcherrimunz. In addi- stunted growth form. Salix brachycarpa is a codominant tion, the GLEES stands differ in their dominance by Salix Figure 14. The Abies lasiocarpa-Picea engelmannii/Salix planifolia (ABLA-PIENISAPL) krummholz association is common in the alpine ranges at the Glacier Lakes Ecosystem Experiments Sites. \ USDA Forest Service Res. Pap. RMRS-RP-1. 1998 planifolia, Salix brachycarpa, and Vacciniunz cespitosunz in An Abies lasiocarpa-Picea engelmannii/Juniperus conzmu- the shrub layer. nis forest type is documented in the literature, but it seems to occur at elevations lower than those at the GLEES and A bies lasiocarpa-Picea engelmannii/Juniperus has forest tree structure (Moir and Ludwig 1979).The cited communis Krummholz type has a richer shrub layer, with Sheperdia canadensis, Synzphoricarpos oreophil us,Mahonia repens, Rosa spp., and R. The Abies Iasiocarpa-Picea engeInzannii/fuzziperus conznzu- wolfii commonly occurring in some or all of the other nis (ABLA-PIEN/JUCO)krummholz association (figure 15) locations. Another low elevation type similar to ABLA- is physiognomically similar to the ABLA-PIEN/SAPL PIEN/ JUCO is the Abies lasiocarpa-Picea engelmannii/ type, but it occurs in drier environments on gentle slopes. Vacciniunz cespitosunz association (Mauk and Henderson Juniperus comnzunis replaces willow as the shrub layer 1984), which has been documented in the Medicine Bow dominant. Salix brachycarpa and Vaccinium cespitosunz are Mountains (Wirsing 1973).Again, there are floristic differ- still important shrub associates; in addition, Ribes ences in both the shrub and grass layers. Since each of the monfigenunz occurs in these stands. Calamagrostis similar types found in the literature have major floristic purpurascens and Carex rossii are important graminoid differences from the GLEES type, this krummholz com- associates, and forbs are a minor component of the com- munity at the GLEES may not have been well described munity. before the current study. Figure 15. The Abies lasiocarpa-Picea engelmannii/Juniperus communis (ABLA-PIENIJUCO) krummholz association is physiog- nomically similar to the ABLA-PIENISAPL type, but it occurs in drier environments on gentle slopes at the Glacier Lakes Ecosys- tem Experiments Sites. \ USDA Forest Service Res. Pap. RMRS-RP-1. 1998 A bies lasiocarpa-Picea engelmannii/Carex rossii documented in the same type elsewhere. Again, we feel Krummholz that the ABLA-PIEN/CARO that we have described at the GLEES differs substantially from what is in the literature A third krummholz association at the GLEES is Abies and may not have been well described before the current lasiocarpa-Picea engelnzannii/Carex rossii (ABLA-PIEN/ study. CARO) (figure 16). These stands occur in the driest of the krummholz environments; on steep, exposed slopes. The Abies lasiocarpa-Picea engelmannii/ understory is usually very sparse and associated species Caltha leptosepala Forest include: Hieraciunz gracile, Juncus drunznzondii, Sibbaldia procun~bens,and Vacciniunz scopariunz. The Picea engelnrannii/Caltha leptosepala (PIEN/CALE) The Abies lasiocarpa-Picea engelnzannii/Carex rossii type forests (figure 17)occur in mesic to wet subalpine environ- documented in the literature (DeVelice et al. 1986) is also ments at the GLEES. They are found in topographic de- found at elevations lower than those at the GLEES. In pressions as patches in a landscape characterized as a addition, there are some floristic differences between the mosaic of forest with sedge meadow (DECE/CANI) open- GLEES sites and other locations, with Paxistinza nzyrsinites ings in the most poorly drained areas. They are also in found in the shrub layer and Osmorhiza chilensis and Ciliaria drainages on streambanks and alluvial terraces. The soils austromontana in the forb layer. There are a number of are sa tura ted most or all of the growing season and surface species found in this type at the GLEES that are not water may be present early in the summer. The most Figure 16. The Abies lasiocarpa-Picea engelmannii/Carex rossii (ABLA-PIENICARO) krummholz association occurs on steep, exposed slopes at the Glacier Lakes Ecosystem Ex'periments Sites. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 important associated species are graminoids and include tributed forest type that is also found on poorly drained Carex illota, Carex nzicroptera, Carex nigricans, and soils (Hess 1981;Wasser and Hess 1982; Hess and Alexander Deschanzpsia cespitosa. Important associated forbs are 1986; Alexander et al. 1986).The ABLA-PIEN/CALE type Saxifraga odontolonza and Trollius laxus. These wet forests at the GLEES seems to occur on even wetter sites. Where are included in the conifer forest landscape type described Calanzagrostis canadensis is dominant in the herb layer, by Simmons (1994). there is a rich forb community with Dodecatheon spp., The Picea engelnzanniilCaltha lepfosepala association is Veratrun~tenuipetalunz, and Thalictrun~occidentale impor- documented in northwestern Wyoming and northeastern tant at other locations but not occurring at the GLEES. In Utah (Steele et al. 1983; Mauk and Henderson 1984). This addition, more species of Carex occur in the wetter GLEES type is very similar to the GLEES association except for the site. presence of Picea pungens in the overstory and Kalmia The Abies Zasiocarpa-Picea engelnzanniilMertensia ciliata polifolia and Phyllodoce empetrifornzis in the shrub and herb association of southern and southwestern Colorado is also layer in the PIEN/CALE stands. In addition, at the GLEES similar but again is found on slightly dryer sites. The Abies lasiocarpa is a codominant and is absent from some of southern association has a much richer forb layer with the other stands, which tend to occur at low elevations. Oxypolis fendleri, Cardarnine cordifolia, Ligularia bigelovii, Peet (1981)describes a Picea-Abies bog forest that is compa- Heracleum sphondylium, Erigeron coulteri, Moneses uniflora, rable to our ABLA-PIEN/CALE type. The Abies lasiocarpa- Polemoniunz pulcherrinzunz, Geraniunz richardsonii, and Picea engeEmannii/Calamagrostiscanadensis is a widely dis- Streptopus fassettii. A third association that is similar to the Figure 17. The Picea engelmannii/Caltha leptosepala (PEINICALE) forests occur in mesic to wet subalpine environments at the Glacier Lakes Ecosystem Experiments Sites. \ USDA Forest Service Res. Pap. RMRS-RP-1. 1998 ABLA-PIEN/CALE type at the GLEES is Abies lasiocarpa- tion of the GLEES can be classified into 4 meadow, 4 Picea engel nzannii/Senecio triangularis (Hess 1981; Peet 1981; shrubland, 3 krummholz, and 2 forest associations that Wasser and Hess 1982; Crouch 1985; Komarkova 1988), differ in floristic composition and community structure, which again occupies drier sites and is more species rich. and that occur repeatedly on the GLEES landscape. Our Species that occur in this type but do not occur at the description of types reflects variability in composition GLEES include: Vacciniunz nzyrtillus in the shrub layer, and structure of nonforested types that is not well docu- Carex disperma, Eleocharis quinqueflora, Scirpus nzicrocarpus, mented in the literature. In addition, 2 krummholz asso- Cardanzine cordifolia, Snrilacina stellata, Streptopus fassettii, ciations, Abies lasiocarpa-Picea engelnzannii/Juniperusconz- Veronica nutans, Moneses uniflora, and Polenzoniunz nzunis krummholz and Abies lasiocarpa-Picea engelnzannii krummholz, were not well documented before this study and the Abies lasiocarpa-Picea engelmannii/Caltha leptosepala Abies lasiocarpa/Vaccinium scoparium Forest wet forests were not well described for this geographic region. Abies lasiocarya/Vacciniunzscopariunz (ABLA-VASC)for- These results provide additional information on the ests cover much of the subalpine forested landscape at the vegetation of subalpine/alpine ecotones in general, and GLEES. These forest occur in a variety of upland sites in specifically they provide a baseline for future studies of mesic to xeric environments. The most mesic of these the vegetation of the GLEES. The plant community classi- forests have a heavy snow accumulation and a prolonged fication provides an organizational framework for design- snow cover. The ground layer vegetation is lush in the ing future studies of the functional characteristics of com- most mesic sites and is dominated by Vacciniunz scopariunz. munities or systems and species-environment relation- In drier environments, the ground layer may be very ships at the GLEES. sparse. Associated species occurring across the full range of these forests include: ~rnicncordifolia, Carex rossii, Erigeron peregrinus, Poa neruosa, and Ribes nzon tigenunz. Picea engelnzannii is present as a codomi- nant in all the GLEES stands. These stands are in- cluded in the coniferous forest landscape type de- Literature Cited scribed by Simmons (1994). This type is one of the most widely distributed in the Alatalo, R.V. 1981. Problems in the measurement of evenness in subalpine-zone and is well documented in the Medicine ecology. Oikos 37:199-204. Bow Mountains (Oosting and Reed 1952; Wirsing 1973; Alexander, R.R., Hoffman, G.R., Wirsing, J.M. 1986. Forest vegetation Wirsing and Alexander 1975; Knight and Thilenius 1975; of the Medicine Bow National Forest in southeastern Wyoming: A Alexander et al. 1986). Our Abies lasiocarpa/Vacciniunz habitat type classification. USDA Forest Service Research Paper RM-271. Rocky Mountain Forest and Range Experiment Station, scopariunz type may include some stands similar to the Fort Collins, Colorado. Allen, R.B., Peet, R.K., Baker, W.L. 1991. Abies lasiocarpa-Picea engelnlannii/Ribes spp. plant associa- Gradient analysis of latitudinal variation in Southern Rocky Moun- tion in the literature (Sheperd 1975; Boyce 1977; Moir and tain forests.Baker, W.L. 1983. Alpine vegetation of the Sangre de Ludwig 1979; Peet 1981; DeVelice et al. 1986; Komarkova Cristo Mountains, New Mexico: gradient analysis, classification, 1988; Allen et al. 1991). 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USDA Forest Strasia, C.A., Thorn, M., Rice, R.W., D.R. Smith. 1970. Grazing habits, Service GTR RM-12/11 pp. diet and performance of sheep on alpine ranges. Journal of Range Youngblood, A.P., Mauk, R.L. 1985. Coniferous forest habitat types of Management 23:201-208. central and southern Utah. USDA Forest Service, Intermountain Ter Braak, C.J.F. 1988. CANOCO - a FORTRAN program for canonical Forest Experiment Station, General Technical Report INT-187, community ordination by [partial] [detrendedl [canonical] corre- 89 PP. spondence analysis, principal components analysis and redun- USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix 1 Vascular plant taxa of the Glacier Lakes Ecosystem Experiments Site. Family Species Equisetaceae Equisetum arvense L. lsoetaceae lsoetes bolanderi Engelm. var. bolanderi Ophioglossaceae Botrychium lunaria (L.) Sw. var. lunaria Polypodiaceae Cryptogramma acrostichoides R. Br. Polypodiaceae Cystopteris fragilis (L.) Bernh . var. fragilis Polypodiaceae Woodsia scopulina D. C. Eat. Selaginellaceae Selaginella densa Ry d b. Cupressaceae Juniperus communis L. var. depressa Pursh Pinaceae Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa Pinaceae Picea engelmannii Parry ex Englem. Pinaceae Pinus contorta Dougl. ex Loud. var. latifolia Engelm. ex Wats. Pinaceae Pinus flexilis James Apiaceae Angelica grayi (Coult. & Rose) Coult. & Rose Apiaceae Conioselinum scopulorum (Gray) Cou It. & Rose Apiaceae Ligusticum porteri Coult. & Rose var. porteri Apiaceae Osmorhiza depauperata Phil. Asteraceae Achillea millefolium L. var. lanulosa (Nutt.) Piper Asteraceae Agoseris aurantiaca (Hook.) Greene Asteraceae Agoseris glauca (Pursh) Raf. var. dasycephala (T. & G.) Jeps. Asteraceae Anaphalis margaritacea (L.) Benth. & Hook. Asteraceae Antenna ria aromatica Evert Asteraceae Antennaria corymbosa E. Nels. Asteraceae Antennaria media Greene Asteraceae Antennaria microphylla Rydb. Asteraceae Antennaria parvifolia Nutt. Asteraceae Antennaria rosea Green e Asteraceae Antennaria umbrinella Ryd b. Asteraceae Arnica cordifolia Hook. Asteraceae Arnica latifolia Bong. Asteraceae Arnica mollis Hook. Asteraceae Arnica ovata Greene Asteraceae Arnica parryi Gray var. parryi Asteraceae Arnica rydbergii Greene Asteraceae Artemisia scopulorum Gray Asteraceae Aster foliaceus Lindl. ex DC. var. apricus Gray Asteraceae Cirsium centaureae (Rydb.) K. Schum. Asteraceae Erigeron compositus Pursh var. discoideus Gray Asteraceae Erigeron melanocephalus (A. Nels.) A. Nels. Asteraceae Erigeron peregrinus (Banks ex Pursh) Greene ssp. callianthemus (Greene) Cronq. Asteraceae Erigeron pinnatisectus (Gray) A. Nels. Asteraceae Erigeron simplex Greene Asteraceae Erigeron ursinus D. C. Eat. Asteraceae Haplopappus pygmaeus (T. & G.) Gray Asteraceae Hieracium gracile Hook. var. gracile Asteraceae Hymenoxys grandiflora (T. & G. ex Gray) Parker Asteraceae Senecio canus Hook. Asteraceae Senecio crassulus Gray Asteraceae Senecio dimorphophyllus Greene var. dimorphophyllus Asteraceae Senecio fremontiiT. & G. var. blitoides (Greene) Cronq. Asteraceae Senecio integerrimus Nutt. var. exaltatus (Nutt.) Cronq. Asteraceae Senecio streptanthifolius Greene USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix 1 (Cont'd.) Family Species Asteraceae Senecio triangularis Hook . Asteraceae Solidago multiradiata Ait var. scopulorum Gray Asteraceae Solidago parryi (Gray) Green e Asteraceae Solidago simplex Ku nt h Asteraceae Taraxacum ceratophorum (Ledeb.) DC. Asteraceae Taraxacum eriophorum R yd b. Asteraceae Taraxacum laevigatum (Wild.) DC. Asteraceae Taraxacum officinale Weber Boraginaceae Eritrichium nanum (Vill.) Schrad. ex Gaudin var. elongatum (Rydb.) Cronq. Boragi naceae Mertensia ciliata (James ex Torr.) G. Don var. ciliata Boraginaceae Mertensia viridis (A. Nels.) A. Nels. Brassicaceae Arabis drummondii Gray Brassicaceae Draba albertina Greene Brassicaceae Draba apiculata C.L. Hitchc. Brassicaceae Draba aurea Vahl ex Hornem. var. aurea Brassicaceae Draba cana Rydb. Brassicaceae Draba crassifolia Grah . Brassicaceae Draba oligosperma Hook. Brassicaceae Rorippa curvipes Greene var: alpha (Wats.) Stuckey Brassicaceae Thlaspi montanum L. var. montanum Callitrichaceae Callitriche palustris L. Campanulaceae Campanula rotundifolia L. Campanulaceae Campanula uniflora L. Caprifoliaceae Sambucus racemosa L. var. microbotrys (Rydb.) Kearney & Peebles Caryoph yllaceae Arenaria congesta N utt. var. congesta Caryophyllaceae Cerastium arvense L. Caryophyllaceae Minuartia obtusiloba (Rydb.) House Caryophyllaceae Minuartia rubella (Wahlenb.) Hiern Caryophyllaceae Paronychia pulvinata Gray Caryophyllaceae Sagina saginoides (L.) Karst. Caryophyllaceae Silene acaulis (L.) Jacq. var. subacaulescens (F. N. Williams) Fern. & St. John Caryophyllaceae Silene drummondii Hook. var. drummondii Caryophyllaceae Silene drummondii Hook. var. striata (Rydb.) Bocq. Caryophyllaceae Silene parryi (Wats.) Hitchc. & Maguire Caryophyllaceae Spergularia rubra (L.) J. & K. Presl Caryophyllaceae Stellaria borealis Bigel. ssp. borealis Caryophyllaceae Stellaria calycantha (Ledeb.) Bong. Caryophyllaceae Stellaria longipes Goldie var. longipes Caryophyllaceae Stellaria monantha H ul t . Caryophyllaceae Stellaria umbellata Turcz. ex Kar. & Kir. Chenopodiaceae Chenopodium atrovirens Rydb. Chenopodiaceae Chenopodium leptophyllum (Moq.) Nutt. ex Wats. Crassulaceae Sedum integrifolium (Raf .) A. Nels. ssp. integrifolium Crassulaceae Sedum lanceolatum Torr. var. lanceolatum Crassulaceae Sedum rhodanthum Gray Cyperaceae Carex albonigra Mack. Cyperaceae Carex aquatilis Wah lenb. var. aquatilis Cyperaceae Carex atrata L. Cyperaceae Carex atrata L. var. chalciolepis (Holm) Kuek. Cyperaceae Carex atrata L. var. erecta Boott Cyperaceae Carex aurea Nutt. Cyperaceae Carex bipartita All. Cyperaceae Carex canescens L. var. cancescens Cyperaceae Carex capilla\ris L. Cyperaceae Carex ebenea Rydb. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix 1 (Cont'd.) Family Species Cyperaceae Carex egglestonii Mack. Cyperaceae Carex foetida All. var. vernacula (Bailey) Kuek. Cyperaceae Carex geyeri Boott Cyperaceae Carex haydeniana Olney Cyperaceae Carex hoodii Boott Cyperaceae Carex illota Bailey Cyperaceae Carex macloviana Urv. Cyperaceae Carex microptera Mack. var. limnophilia (F.J. Herm.) Dorn Cyperaceae Carex microptera Mack. var. microptera Cyperaceae Carex nardina Fries var. hepburnii (Boott) Kukenth. Cyperaceae Carex nelsonii Mack. Cyperaceae Carex nigricans C. A. Mey. Cyperaceae Carex norvegica Retz. var. stevenii (Holm) Dorn Cyperaceae Carex nova Bailey var. nova Cyperaceae Carex pachystacyhya Cham. ex Steud. Cyperaceae Carex phaeocephala Piper Cyperaceae Carex praeceptorum Mack. Cyperaceae Carex pyrenaica Wahlenb. Cyperaceae Carex raynoldsii Dewey Cyperaceae Carex rossii Boott Cyperaceae Carex rostrata Stokes ex With. var. rostrata Cyperaceae Carex rupestris All. Cyperaceae Carex saxatilis L. var. major Olney Cyperaceae Carex scopulorum var. bracteosa C yperaceae Carex scopulorum Holm var. scopulorum Cyperaceae Carex vesicaria L. var. vesicaria Cyperaceae Eleocharis pauciflora (Lightf .) Link Ericaceae Arctostaphylos uva-ursi (L.) Spreng. ssp. uva-ursi var. stipitata (Packer ex Denford) Ericaceae Arctostaphylos uva-ursi (L.) Spreng. ssp. uva-ursi var. uva-ursi Ericaceae Gaultheria humifusa (Grah.) Rydb. Ericaceae Kalmia microphylla (Hook.) Heller var. microphylla Ericaceae Monotropa hypopithys L. Ericaceae Orthilia secunda (L.) House Ericaceae Pyrola minor L. Ericaceae Vaccinium cespitosum Michx. Ericaceae Vaccinium scoparium Leib. ex Cov. Fabaceae Astragalus alpinus L. var. alpinus Fabaceae Lupinus argenteus Pursh var. argenteus Fabaceae Oxytropis campestris var. (L.) DC. var. gracilis (A. Nels.) Barneby Fabaceae Trifolium dasyphyllum T. & G. Fabaceae Trifolium parryi Gray Gentianaceae Gentiana algida Pall. Gentianaceae Gentiana parryi Engelm. Gentianaceae Gentianella amarella (L.) Boerner var. amarella Gentianaceae Gentianopsis detonsa (Rottb.) Ma var. elegans (A. Nels.) N. Holmgren Gentianaceae Swertia perennis L. Grossulariaceae Ribes lacustre (Pers.) Poir. Grossulariaceae Ribes montigenum McClat. Hydrocharitaceae Elodea canadensis Michx. Hydrophyllaceae Phacelia sericea (Grah. ex Hook.) Gray var. ciliosa Rydb. Hydrophyllaceae Phacelia sericea (Grah. ex Hook.) Gray var. sericea l ridaceae Iris missouriensis Nutt. Juncaceae Juncus castaneus Sm. Juncaceae Juncus confusus Gov. USDA Forest Service Res. Pap. RMRS-RP-1 . 1998 Appendix 1 (Cont'd.) - -- Family Species Juncaceae Juncus drummondii E. Mey Juncaceae Juncus drummondii E. Mey . var. drummondii Juncaceae Juncus drummondii E. Mey. var. subtriflorus (E. Mey.) C. L. Hitchc. Ju ncaceae Juncus hallii Engelm. Juncaceae Juncus mertensianus Bong. Juncaceae Juncus parryi Engelm. Ju ncaceae Juncus triglumis L. var. albescens Lange Juncaceae Luzula multiflora (Ehrh .) Lej. Juncaceae Luzula parviflora (Ehrh.) Desv. Juncaceae Luzula spicata (L.) DC. Liliaceae Allium brevistylum Wats. Liliaceae Erythronium grandiflorum Pursh var. grandiflorum Liliaceae Streptopus amplexifolius (L.) DC. Liliaceae Zigadenus elegans Pursh Onagraceae Epilobium anagallidifolium Lam . Onagraceae Epilobium angustifolium L. var. angustifolium Onagraceae Epilobium angustifolium L. var. canescens Wood Onagraceae Epilobium ciliatum Raf . var. ciliatum Onagraceae Epilobium ciliatum var. glandulosum Onagraceae Epilobium clavatum Trel. Onagraceae Epilobium halleanum Hausskn. Onagraceae Epilobium hornemannii Reichenb. ssp. hornemannii Onagraceae Epilobium lactiflorum Hauss kn. Onagraceae Epilobium saximontanum Hausskn . Onagraceae Gayophytum decipiens Lewis & Szweyk. Onagraceae Gayophytum diffusum T. & G. var. strictipes (Hook) Dorn Onagraceae Gayophytum racemosum T. & G. Orchidaceae Platanthera dilatata (Pursh) Lindl. ex Beck var. albiflora (Cham.) Ledeb. Orchidaceae Platanthera dilatata (Pursh) Lindl. ex Beck var. dilatata Orobanc haceae Orobanche uniflora L. var. occidentalis (Greene) Taylor & MacBryde Plantaginaceae Plantago tweedyi Gray Poaceae Agropyron subsecundum (Link) Hitchc. Poaceae Agrostis humilis Vasey Poaceae Agrostis scabra Willd. var. scabra Poaceae Agrostis thurberiana Hitc hc. Poaceae Agrostis variabilis R yd b. Poaceae Bromus ciliatus L. Poaceae Bromus inermis Leyss. var. inermis Poaceae Calamagrostis canadensis (Michx.) Beauv. Poaceae Calamagrostis purpurascens R. Br. var. purpurascens Poaceae Danthonia intermedia Vasey Poaceae Deschampsia atropurpurea (Wahlenb.) Scheele var. latifolia (Hook.) Scribn. ex Macoun Poaceae Deschampsia cespitosa (L.) Beauv. var. cespitosa Poaceae Elymus glaucus Buckl. var. glaucus Poaceae Elymus scribneri (Vasey) Jones Poaceae Elymus trachycaulus (Link) Gould ex Shinners var. trachycaulus Poaceae Festuca brachyphylla Schult. & Schult. ssp. coloradensis Frederiksen Poaceae Festuca saximontana Rydb. var. saximontana Poaceae Hierochloa odorata (L.) Beauv. Poaceae Oryzopsis exigua T hu r b. Poaceae Phleum alpinum L. var. alpinum Poaceae Poa alpina L. Poaceae Poa arctica R. Br. var. grayana (Vasey) Dorn Poaceae Poa compressa L. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix 1 (Cont'd.) Family Species Poaceae Poa cusickii Vasey var. epilis (Scribn.) C. L. Hitchc. Poaceae Poa fendleriana (Steud.) Vasey Poaceae Poa interior Rydb. Poaceae Poa leptocoma Trin. Poaceae Poa nervosa (Hook.) Vasey var. wheeleri (Vasey) C. L. Hitchc. Poaceae Poa pattersonii Vasey Poaceae Poa pratensis L. Poaceae Poa reflexa Vasey & Scribn. Poaceae Poa rupicola Nash ex Rydb. Poaceae Poa secunda Presl var. elongata (Vasey) Dorn Poaceae Poa secunda Presl var. incurva (Scribn. & Williams ex Scribn.) Beetle Poaceae Stipa lettermanii Vasey Poaceae Torreyochloa pallida (Torr.) Church var. pauciflora (J. Presl) J. Davis Poaceae Trisetum spicatum (L.) Richt. Poaceae Trisetum wolfii Vase y Polemoniaceae Collomia linearis Nutt . Polemoniaceae Phlox multiflora A. Nels. Polemoniaceae Phlox pulvinata (Wherry) Cronq. Polemoniaceae Polemonium brandegei (Gray) Greene Polemoniaceae Polemonium viscosum Nutt. Polygonaceae Eriogonum umbellatum Torr. var. majus Hook. Polygonaceae Oxyria digyna (L.) Hill Polygonaceae Polygonum bistortoides Pursh Polygonaceae Polygonum douglasii var. douglasii Polygonaceae Polygonum sawatchense Sm. Polygonaceae Polygonum viviparum L. Polygonaceae Rumex densiflorus Osterh. Polygonaceae Rumex utahenses Rech. f. Portulacaceae Claytonia lanceolata Purs h var. lanceolata Portulacaceae Lewisia pygmaea (Gray) Robins. var. pygmaea Portulacaceae Lewisia triphylla (Wats.) Robins. Primulaceae Androsace septentrionalis L. var. subulifera Gray Primulaceae Primula parryi Gray Ranunculaceae Aconitum columbianum Nutt. var. columbianum Ranunculaceae Anemone multifida Poir. var. multifida Ranunculaceae Anemone patens L. var. multifida Pritz. Ranuncuiaceae Aquilegia coerulea James var. coerulea Ranunculaceae Caltha leptosepala DC. ssp. leptosepala var. leptosepala Ranunculaceae Delphinium barbeyi (Huth) Huth Ranunculaceae Ranunculus alismifolius Geyer ex Benth. var. montanus Wats. Ranunculaceae Ranunculus eschscholtzii Sc hlect. var. eschscholtzii Ranunculaceae Ranunculus flammula L. Ranunculaceae Ranunculus flammula L. var. filiformis (Michx.) Hook. Ranunculaceae Ranunculus inamoenus Greene var. alpeophilus (A. Nels.) Benson Ranunculaceae Ranunculus uncinatus D. Don ex G. Don Ranunculaceae Trollius laxus Salis b. var. albiflorus Gray Rosaceae Geum rossii (R. Br.) Ser. var. turbinatum (Rydb.) C. L. Hitchc. Rosaceae Potentilla concinna Richards. var. concinna Rosaceae Potentilla diversifolia Leh m . var. diversifolia Rosaceae Poteneilla fruticosa L. Rosaceae Potentilla gracilis Dougl. ex Hook. var. nuttallii (Lehm.) Sheld Rosaceae Potentilla gracilis Dougl. ex Hook. var. pulcherrima (Lehm.) Fern Rosaceae Potentilla hookeriana Lehm. , Rosaceae Potentilla nivea 4. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix I (Cont'd.) Family Species Rosaceae Potentilla plattensis Nutt. Rosaceae Potentilla rubricaulis Leh m. Rosaceae Rubus idaeus L. var. aculeatissimus Regel & Tiling Rosaceae Sibbaldia procumbens L. Rubiaceae Galium trifidum L. var. trifidum Salicaceae Populus tremuloides Mic hx. Salicaceae Salix arctica Pall. var. petraea Anderss. Salicaceae Salix bebbiana Sarg. var. bebbiana Salicaceae Salix brachycarpa Nutt. var. brachycarpa Salicaceae Salix cascadensis Ckl I. Salicaceae Salix glauca L. var. villosa (Hook.) Anderss. Salicaceae Salix lasiandra Benth. var. caudata (Nutt.) Sudw. Salicaceae Salix planifolia Pu rs h var . planifolia Saxifragaceae Heuchera parvifolia Nutt. ex T. & G. Saxifragaceae Lithophragma glabrum Nutt. var. ramulosum (Suksd.) Boivin Saxifragaceae Mitella pentandra Hook. Saxif ragaceae Parnassia fimbriata Konig var. fimbriata Saxifragaceae Saxifraga odontoloma Pi per Saxifragaceae Saxifraga rhomboidea Greene Saxif ragaceae Saxifraga rivularis Greene Saxifragaceae Saxifraga rivularis L. ssp. hyperborea (R. Br.) Dorn var. debilis (Engelm ex Gray) Dorn Scrophulariaceae Castilleja rhexifolia Ry d b. Scrophulariaceae Castilleja sulphurea Rydb. Scrophulariaceae Chionophila jamesii Benth. Scrophulariaceae Pedicularis bracteosa Benth. var. paysoniana (Penn.) Cronq. Scrophulariaceae Pedicularis groenlandica Retz. Scrophulariaceae Pedicularis parryi Gray ssp. parryi Scrophulariaceae Pedicularis racemosa Dougl. ex Hook. var. alba (Penn.) Cronq. Scrophulariaceae Penstemon procerus Dougl. ex Grah. var. procerus Scrophulariaceae Penstemon rydbergii A. Nels. var. rydbergii Scrophulariaceae Penstemon whippleanus Gray Scrophulariaceae Veronica serpyllifolia L. var. humifusa (Dickson) Vahl Scrophulariaceae Veronica wormskjoldii R. & S. Violaceae Viola adunca Sm. Violaceae Viola palustris L. Violaceae Viola praemorsa Dougl. ex Lindl. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix 2 Vascular plant taxa of the Glacier Lakes Ecosystem Experiments Site ranked by presence (the number of stands where the species occurred) and percent occurrence (the proportion of stands where present). Taxa present at the GLEES but not occurring in the sample stands are not included. Species occurrence Presence YO Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa 83 Erigeron peregrinus (Banks ex Pursh) Greene spp. callianthemus (Greene) Cronq. 83 Hieracium gracile Hook. var. gracile Picea engelmannii Parry ex Englem. Vaccinium scoparium Leib. ex Cov. Potentilla diversifolia Leh m . var. diversifolia Carex rossii Boott Polygonum bistortoides Purs h Sibbaldia procumbens L. Senecio dimorphophyllus Greene var. dimorphophyllus Achillea millefolium L. var. lanulosa (Nutt.) Piper Juncus drummondii E. Mey Phleum alpinum L. var. alpinum Penstemon whippleanus Gray Trisetum spicatum (L.) Richt. Lewisia pygmaea (Gray) Robins. var. pygmaea Erythronium grandiflorum P urs h var. grandiflorum Selaginella densa Ryd b. Veronica wormskjoldii R . & S. Deschampsia cespitosa (L.) Beauv. var. cespitosa Ribes montigenum McClat. Senecio crassulus Gray Antennaria umbrinella Rydb. Artemisia scopulorum Gray Sedum lanceolatum Torr. var. lanceolatum A rnica cordifolia Hook . Caltha leptosepala DC. ssp. leptosepala var. leptosepala Minuartia obtusiloba (R ydb.) House Poa nervosa (Hook.) Vasey var. wheeleri (Vasey) C. L. Hitchc. Poa reflexa Vasey & Scribn. Poa cusickii Vasey var. epilis (Scribn.) C. L. Hitchc. Geum rossii (R. Br.) Ser. var. turbinatum (Rydb.) C. L. Hitchc. Thlaspi montanum L. var. montanum Carex nigricans C. A. Mey. Salix planifolia Pursh var. planifolia Pedicularis racemosa Dougl. ex Hook. var. alba (Penn.) Cronq. Sedum rhodanthum Gray Vaccinium cespitosum Mic hx. Danthonia intermedia Vasey Erigeron pinnatisectus (Gray) A. Nels. Epilobium angustifolium L. var. canescens Wood Agoseris glauca (Pursh) Raf. var. dasycephala (T. & G.) Jeps. Festuca brachyphylla Schult. & Schult. spp. coloradensis Frederiksen Pedicularis bracteosa Benth. var. paysoniana (Penn .) Cronq. Pedicularis groenlandica Retz. Arenaria congesta Nutt. var. congesta Erigeron melanocephalus (A. Nels.) A. Nels. \ USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix 2 (Cont'd.) Species occurrence Presence YO Arnica mollis Hook. Luzula spicata (L.) DC. Poa alpina L. Agrostis thurberiana Hitchc. Arnica latifolia Bong. Carex aquatilis Wahlenb. var. aquatilis Viola adunca Sm. Aquilegia coerulea James var. coerulea Gentianella amarella (L.) Boerner var. amarella Minuartia rubella (Wahlenb.) Hiern Trifolium parryi Gray Conioselinum scopulorum (Gray) Cou It. & Rose Juniperus communis L. var. depressa Pursh Solidago simplex K un t h Carex microptera Mack. var. microptera Phlox pulvinata (Wherry) Cronq. Arabis drummondii Gray Carex rupestris A lI. Pedicularis parryi G ray ssp. parryi Polemonium viscosum N utt. Antennaria media Greene Epilobium hornemannii Reichenb. spp. hornemannii Juncus mertensianus Bong. Luzula parviflora (Ehr h .) Desv. Salix brachycarpa Nutt. var. brachycarpa Angelica grayi (Coult. & Rose) Coult. & Rose Erigeron simplex Green e Juncus parryi Engelm. Poa fendleriana (Steud.) Vasey Potentilla nivea L. Ranunculus alismifolius Geyer ex Benth. var. montanus Wats. Calamagrostis purpurascens R. Br. var. purpurascens Cerastium arvense L. Kalmia microphylla (Hook.) Heller var. microphylla Senecio triangularis Hook. Silene acaulis (L.) Jacq. var. subacaulescens (F. N. Williams) Fern. & St. John Taraxacum officinale Weber Agrostis humilis Vasey Calamagrostis canadensis (Michx.) Beauv. Carex illota Bailey Carex scopulorum Holm var. scopulorum Draba oligosperma Hook. Eritrichium nanum (Vill.) Schrad. ex Gaudin var. elongatum (Rydb.) Cronq. Festuca saximontana Ryd b. var. saximontana Arctostaphylos uva-ursi (L.) Spreng. spp. uva-ursi var. stipitata (Packer ex Denford) Dorn Castilleja rhexifolia Rydb. Mertensia ciliata (James ex Torr.) G. Don var. ciliata Paronychia pulvinata Gray Trifolium dasyphyllum T. & G. Androsace septentrionalis L. var. subulifera Gray Arnica rydbergii Green e x Carex ebenea Rydb. USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix 2 (Cont'd.) Species occurrence Presence O/O Deschampsia atropurpurea (Wa hlenb.) Scheele var. latifolia (Hook.) Scribn. ex Macoun Draba aurea Vahl ex Hornem. var. aurea Hymenoxys grandiflora (T. & G. ex Gray) Parker Platanthera dilatata (Pursh) Lindl. ex Beck var. albiflora (Cham.) Ledeb. Poa interior Rydb. Poa pattersonii Vasey Poa secunda Presl var. elongata (Vasey) Dorn Polygonum viviparum L. Saxifraga odontoloma Piper Saxifraga rhomboidea Greene Stellaria umbellata Turcz. ex Kar. & Kir. Trisetum wolfii Vasey Antennaria rosea Greene Aster foliaceus Lindl. ex DC. var. apricus Gray Epilobium halleanum Hausskn. Haplopappus pygmaeus (T. & G.) Gray Plantago tweedyi Gray Poa leptocoma Trin. Ranunculus eschscholtzii Sc hlect. var. eschscholtzii Ranunculus inamoenus Greene var. alpeophilus (A. Nels.) Benson Agrostis variabilis Rydb. Carex atrata L. var. erecta Boott Carex foetida All. var. vernacula (Bailey) Kuek. Carex pyrenaica Wahlen b. Carex scopulorum var. bracteosa Elymus scribneri (Vasey) Jones Erigeron ursinus D. C. Eat. Gentiana algida Pal I. Mitella pentandra Hook. Pinus flexilis James Salix cascadensis C klI. Solidago multiradiata Ai t var. scopulorum Gray Carex albonigra Mack. Carex haydeniana Olney Cystopteris fragilis (L.) Bern h. var. fragilis Elymus trachycaulus (Link) Gould ex Shinners var. trachycaulus Mertensia viridis (A. Nels.) A. Nels. Astragalus alpinus L. var. alpinus Carex atrata L. var. chalciolepis (Holm) Kuek. Carex canescens L. var. cancescens Carex pachystacyhya Cham. ex Steud. Carex phaeocephala Piper Chionophila jamesii Benth. Draba crassifolia Grah . Eleocharis pauciflora (Lightf .) Link Epilobium lactiflorum Hausskn. Gentiana parryi Engelm. Oxyria digyna (L.) Hill Phacelia sericea (Grah. ex Hook.) Gray var. sericea Pinus contorta Dougl. ex Loud. var. latifolia Engelm. ex Wats. Poa rupicola Nash ex Rydb. Rumex densiflorus Osterh . Sagina saginoides (L.) Karst. k USDA Forest Service Res. Pap. RMRS-RP-1. 1998 Appendix 2 (Cont'd.) Species occurrence Presence O/O Senecio fremontii T. & G. var. blitoides (Greene) Cronq. Stellaria borealis Bigel. spp. borealis Swertia perennis L. Viola praemorsa Dougl. ex Lindl. Agoseris aurantiaca (Hook.) Greene Agrostis scabra Willd. var. scabra Antennaria microphylla Rydb. Botrychium lunaria (L.) Sw. var. lunaria Callitriche palustris L. Carex hoodii Boott Carex microptera Mack. var. limnophilia (F.J. Herm.) Dorn Carex nelsonii Mack. Carex nova Bailey var. nova Carex praeceptorum Mack. Carex raynoldsii Dewey Carex vesicaria L. var. vesicaria Draba albertina Greene Elymus glaucus Buckl. var. glaucus Epilobium ciliatum Raf. var. ciliatum Epilobium cla vatum Trel. Epilobium saximontanum Hausskn. Equisetum arvense L. Gaultheria humifusa (Grah.) Rydb. Hierochloa odorata (L.) Beauv. Juncus con fusus Cov. Lupinus argenteus Pursh var. argenteus Monotropa hypopithys L. Oryzopsis exigua T h urb. Osmorhiza depauperata Phil. Oxytropis campestris var. (L.) DC. var. gracilis (A. Nels.) Barneby Parnassia fimbriata Konig var. fimbriata Poa compressa L. Polygonum douglasii va r. douglasii Populus tremuloides M ic h x. Pyrola minor L. Ranunculus flammula L. var. filiformis (Mic hx.) Hook. Rorippa curvipes Greene var. alpina (Wats.) Stuckey Sambucus racemosa L. var. microbotrys (Ryd b.) Kearney & Peebles Saxifraga rivularis G reene Senecio streptanthifolius Greene Silene drummondii Hook. var. striata (Rydb.) Bocq. Solidago parry; (Gray) Greene Stellaria calycantha (Ledeb.) Bong. Stellaria longipes Goldie var. longipes Stellaria monantha Hult. Streptopus amplexifolius (L.) DC. Taraxacum ceratophorum (Ledeb.) DC. Taraxacum laevigatum (Willd.) DC. Woodsia scopulina D. C. Eat. Zigadenus elegans Pursh USDA Forest Service Res. Pap. RMRS-RP-1. 1998 ROCKY9 MOUNTAIN The Rocky Mountain Research Station develops scientific information and technology to improve management, protection, and use of forests and rangelands. 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